QL
461
.A373X
ENT
MEMOIRS
OF THE
INTOMOLOGICAL SOCIETY
NUMBER 24
THE HIGHER CLASSIFICATION,
PHYLOGENY AND ZOOGEOGRAPHY
OF THE SATYRIDAE (LEPIDOPTERA)
BY
LEE D. MILLER
BL1SHED BY THE AMEBJ INTOMOLOGICAL SOCIETY
AT THE ACADEMY OF NATURAL SCIENCES
PHILADELPHIA
1968
MEMOIRS
OF THE
AMERICAN ENTOMOLOGICAL SOCIETY
NUMBER 24
THE HIGHER CLASSIFICATION,
PHYLOGENY AND ZOOGEOGRAPHY
OF THE SATYRIDAE (LEPIDOPTERA)
BY
LEE D. MILLER
PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY
AT THE ACADEMY OF NATURAL SCIENCES
PHILADELPHIA
1968
Selwyn S. Roback
Editor
(Issued October 22, 1968)
Printed in the United States of America
To the memory of Dr. Richard M. Fox
— Teacher, inspiration, colleague, friend.
He is and will be missed.
Preface
No work is ever truly done alone: this paper is ample proof of
the statement. I wish to take this opportunity to thank the many
people and organizations who have helped to make it a reality.
The research was supported in part by National Science Founda-
tion grant number GB-2928 (R. M. Fox, Principal Investigator).
The National Science Foundation, through Office of Science Infor-
mation Services, made possible the publication of this work by grant
number GN-652.
Thanks are due the following individuals and institutions for their
aid, loan of materials, use of facilities and general encouragement:
Carnegie Museum, particularly Dr. M. Graham Netting, Director,
Drs. Richard M. Fox and George E. Wallace and Mr. Harry K.
Clench of the Section of Insects and Spiders; the Department of En-
tomology, British Museum (Natural History), especially Messrs. J. P.
Doncaster, Keeper of Entomology, and N. D. Riley, P. E. S. Whal-
ley, T. G. Howarth and M. Clifton of the Lepidoptera section; and
the Departments of Biology of the University of Pittsburgh and The
Catholic University of America. Without their cooperation this pro-
ject could not have been completed.
The following individuals read the original manuscript which was
submitted in quite different form as a dissertation at the University of
Pittsburgh: Drs. Fox, Peter Gray, Eliot B. Spiess, Malcolm T. Jollie
and Kenneth W. Cummins, Department of Biology, University of
Pittsburgh, C. C. Li, Department of Biostatistics, Graduate School of
Public Health. University of Pittsburgh, Craig C. Black and Mary
Dawson, Section of Vertebrate Paleontology, Carnegie Museum. All
were available for consultation and deserve great thanks.
Mr. Clifton read the part of the manuscript dealing with the
Haeterinae and made the necessary revisions on it based upon his
forthcoming revision. Dr. Norman Tindale of the South Australian
Museum read and commented upon the zoogeographic analyses, espe-
cially those dealing with Australia. Dr. F. Martin Brown and Mr.
Harry K. Clench read all or part of the manuscript and commented
upon it. Particular thanks, however, are due Dr. Fox, who read and
carefully critiqued the entire manuscript. He offered countless valu-
able suggestions, most of which have been incorporated into the manu-
script.
Preparation of the manuscript and the figures has been aided
greatly through the activities of two of my graduate students, Misses
Jacqueline Y. Sessi and Dorothy A. Shea.
Finally, thanks are due the many unnamed friends and colleagues
who, knowingly or not, have contributed ideas and moral support for
this undertaking.
Table of Contents
Page
INTRODUCTION 1
The Problem 1
Historical Sketch 1
Material Studied and Methods Employed 4
COMPARATIVE MORPHOLOGY 7
Antennae 7
Head 7
Labial Palpi 7
Thorax 8
Forelegs 8
Male forelegs 8
Female forelegs 9
Walking Legs 10
Wings 10
Forewings 1 1
Hindwings 12
Androconial patches 13
Male Genitalia 14
SYSTEMATIC REVISION OF THE GENERA OF THE SATYRIDAE 14
Family Satyridae Boisduval 15
Key to the Subfamilies of the Satyridae 18
Subfamily Haeterinae Herrich-Schaffer 19
Tribe Haeterini Herrich-Schaffer 20
Genera Included in the Haeterini 22
Subfamily Brassolinae Boisduval 23
Tribe Brassolini Boisduval 23
Genera Included in the Brassolini 27
Subfamily Biinae Herrich-Schaffer 28
Key to the Tribes of the Biinae 29
Tribe Antirrhini, new tribe 29
Genera Included in the Antirrhini 31
Tribe Biini Herrich-Schaffer 32
Genus Included in the Biini 34
Tribe Melanitini, new tribe 34
Genera Included in the Melanitini 36
Genus Manataria Kirby 36
Subfamily Elymniinae Herrich-Schaffer 38
Key to the Tribes of the Elymniinae 40
Tribe Lethini Clark 40
Genera Included in the Lethini 47
MEM. AMER. ENT. SOC, 24
Tribe Zetherini, new tribe 52
Genera Included in the Zetherini 53
Tribe Elymniini Herrich-Schaffer 54
Genera Included in the Elymniini 56
Tribe Mycalesini, new tribe 57
Genera Included in the Mycalesini 60
Subfamily Eritinae, new subfamily 64
Tribe Eritini, new tribe 64
Genera Included in the Eritini 66
Subfamily Ragadiinae Herrich-Schaffer 67
Tribe Ragadiini Herrich-Schaffer 67
Genera Included in the Ragadiini 69
Subfamily Satyrinae Boisduval 69
Key to the Tribes of the Satyrinae 73
Tribe Hypocystini, new tribe 74
Genera Included in the Hypocystini 80
Tribe Ypthimini, new tribe 81
Genera Included in the Ypthimini 85
Genus Palaeonympha Butler 87
Tribe Euptychiini, new tribe 89
Genera Included in the Euptychiini 92
Tribe Coenonymphini, new tribe 95
Genera Included in the Coenonymphini 98
Tribe Maniolini Hampson 98
Genera Included in the Maniolini 101
Tribe Erebiini Tutt 101
Genera Included in the Erebiini 104
Tribe Dirini, new tribe 105
Genera Included in the Dirini 108
Tribe Pronophilini Clark 108
Genera Included in the Pronophilini 114
Tribe Satyrini Boisduval 119
Genera Included in the Satyrini 123
Tribe Melanargiini Verity 124
Genera Included in the Melanargiini 126
Genera of Uncertain Position 127
Genus Pamperis Heimlich 127
Genus Setodocis Billberg 128
u
THE EVOLUTION AND ZOOGEOGRAPHY OF THE SATYRIDAE 128
Origin of the Satyridae 131
The Later History of the Satyridae 134
Derivation of the Satyrid Faunae of the World 142
The Neotropics 145
The Nearctic 145
The Palearctic 147
The Indo-Malayan Region 148
The Australian Region, including the Pacific Islands 149
The Ethiopian Region 150
Madagascar 152
BIBLIOGRAPHY 153
INDEX 164
MEM. AMER. ENT. SOC, 24
iii
Memoirs
of THE
American Entomological Society
Number 24
THE HIGHER CLASSIFICATION,
PHYLOGENY AND zoogeography OF
THE SATYRIDAE (LEPIDOPTERA)
By
Lee D. Miller
Introduction
The Problem. — The objective of this study, briefly, is the anal-
ysis, both in space and, where possible, in time, of the evolution, phy-
logeny and zoogeography of the higher taxa of the Satyridae. The
existing studies on the family have been generally regional, and little
progress has been made on interrelating the faunae of the various re-
gions of the world. Furthermore, these studies were not based on
modern, multi-variant taxonomic principles; hence, as will be shown,
some fundamental errors in relationships have been perpetrated. Since
no adequate taxonomic revision of the higher categories of the Saty-
ridae is available, it has been necessary to complete one as a basis for
the evolutionary and zoogeographic discussions. Using morphologi-
cal and distributional data as bases, a scheme for the evolution and
zoogeography of the Satyridae is proposed, and finally a phylogeny of
the family is projected.
Historical Sketch. — Linne considered all butterflies congen-
eric. In the Tenth Edition of his Sy sterna Naturae (1758), the start-
ing point of zoological nomenclature, he published descriptions of a
MEM. AMER. ENT. SOC, 24
THE SATYRIDAE
number of satyrids, placing them with all other butterflies in the genus
Papilio. The first nomenclatorially valid satyrid genus, Maniola, was
erected by Schrank (1801) in his Fauna Boica. Fabricius (1807)
described several more genera in Illiger's rare Magazin fur Inseckt-
enkunde. Latreille (1810) proposed Satyrus in his Considerations
generates sur Yordre natural des animaux . . . crustaces . . . arach-
nides . . . et insectes . . . Hiibner, between the years 1806 and
1819 (the latter date is bibliographically indeterminate and is brack-
eted in all succeeding references), published many satyrid generic
names in the Sammlung Exotischer Schmetterlinge and the Vcrzeich-
niss bekannter Schmettlinge [sic]. Several other authors, notably
Billberg (1820) and Meigen (1829), proposed early generic names
for satyrids, but it was not until 1836 that the family was described,
as "Satyrides", by Boisduval. That author also first proposed the
Brassolinae, as "Brassolides".
The first comprehensive work devoted to the butterflies of the
world in which the treatment was anything like a modern one was
The genera of diurnal Lepidoptera by E. Doubleday, J. O. Westwood
and W. C. Hewitson which appeared in two volumes between 1846
and 1852. In this work the Satyridae were defined, the brassolines
delimited (as a subfamily of the Morphidae) and many generic names
were added to the Satyridae. The practice of separating the brasso-
lines from the rest of the satyrids became firmly established with this
publication. The majority of the satyrid generic names were de-
scribed in this work between 1849 and 1851, after Doubleday's death,
and are usually considered to have been authored by Westwood.
The authors always referred to this work as "Doubleday, Westwood
and Hewitson", even though Hewitsons chief role was as illustrator;
but Brown (1941) holds that text descriptions must be credited to
either Doubleday or Westwood alone, while names first proposed in
plates (the text and the plates were often separately published) should
be credited to Doubleday and Hewitson or to Westwood and Hewit-
son. Accordingly, in those instances where the text appeared simul-
taneously with, or prior to, the plates, I am crediting authorship to
either Doubleday or Westwood, but if the plates appeared before the
text descriptions, credit must be given to Doubleday and Hewitson or
to Westwood and Hewitson.
The first attempt at a higher classification of the satyrids ( actually,
LEE D. MILLER 3
of all the butterflies) was that of Herrich-Schaffer (1864). For the
first time the satyrids were split beyond the Satyrinae and Brassolinae,
as "Satyrina" and "Brassolina". The Biinae ("Biina"), Haeterinae
("Hetaerina" [sic]), Ragadiinae ("Ragadiina") and Elymniinae
("Elymniina") were established. This work has been too long over-
looked by systematists — all but one of the presently recognized saty-
rid subfamilies were defined in it. Rober (1892), in the second vol-
ume of Staudinger and Schatz' Exotische Schmetterlinge, divided the
satyrid genera into six groups with a residue of indeterminate genera.
Rober's groups generally correspond to major divisions of the Saty-
ridae, but they were not nomenclatorially designated. Moore in the
first and second volumes of his Lepidoptera Indica, published between
1890 and 1893, proposed a great many generic names but considered
these genera to be apportioned into only the Satyrinae and Elym-
niinae. These same divisions are to be found in Gaede ( 193 1 ) . The
Rober scheme was followed with little modification by Fruhstorfer,
Weymer and Aurivillius in their respective sections of Seitz' monu-
mental Die Grossschmetterlinge der Erde.
More recently Clark (1947) subdivided the satyrids, exclusive of
the brassolines, into the Satyrinae, Enodiinae, Pronophilinae, Elym-
niinae and Pierellinae (a synonym of Haeterinae). In 1948 he
emended Enodiinae to Lethinae and divided the brassolines, as a sep-
arate family, into the Brassolinae, Caliginae and Biinae (this last not
properly a brassoline). In neither paper did Clark give definitive
reasons for his classification, hence he has been criticized by such
authors as Ehrlich (1958) who proposed a higher classification of
the "true butterflies" (Papilionoidea), retaining the separation of the
brassolines and the satyrids and placing the former with the Mor-
phidae.
Table 1 attempts to relate the classifications of Herrich-Schaffer
(1864) and Clark (1947, 1948) with that proposed in the present
paper. In many instances particular genera were not apportioned by
either Herrich-Schaffer or Clark, and those authors' intentions can
only be guessed.
With the exception of Clark (1947, 1948) and Ehrlich (1958),
no modern author has attempted to place the satyrids in their proper
evolutionary positions, and there has been no zoogeographic analysis
of this family. Schwanwitsch (1924), however, relied heavily on
MEM. AMER. ENT. SOC, 24
THE SATYRIDAE
Table 1
A comparison of the present subfamilial classification of the Satyridae with
those proposed by Herrich-Schaffer (1864) and Clark (1947, 1948).
Present Herrich-Schaffer's Clark's
classification classification classification
Haeterinae Hetaerina (sic) Pierellinae
Brassolinae Brassolina Brassolinae
Caliginae
Biinae Biina Biinae (of Brassolidae)
Satyrina (part) Satyrinae (part)
Elymniinae Elymniina Elymniinae
Satyrina (part) Enodiinae
(=Lethinae)
Eritinae Satyrina Satyrinae
Ragadiinae Ragadiina Satyrinae
Satyrinae Satyrina Satyrinae (part)
Pronophilinae
satyrids in his study of the evolutionary trends toward the develop-
ment of the "basic nymphalid pattern". This work was followed by
a number of other papers concerning pattern modifications, two of
which are of interest here: in 1925 he published an analysis of the
Pierella-type pattern, a paper which has since become a classic in the
explanation of the morphological migration of pattern elements; and
in 1931 he analyzed the highly modified Melanargia-type pattern.
Both papers were significant, but the one explaining "pierellization"
has had the most far-reaching implications.
Material Studied and Methods Employed. — Since there are
between 2500 and 3000 species in the Satyridae, it would be im-
practical to attempt a complete revision of all the species in a single
work. The number of described genera, on the other hand, is man-
ageable — fewer than 400.
A fair axiom in systematics may be stated more or less as fol-
lows: systematic decisions at one taxonomic level should be reached
by examination of most (preferably all) of the entities at the next
lower major taxonomic level. In other words, if one is revising a
genus, one should see representatives of the species and subspecies
included in it to make meaningful taxonomic judgments at the spe-
cific and generic levels. In the present study examples of the already-
named genera were examined, hence, taxonomic judgments are made
at the tribal level, and no synonymization of nomenclatorially valid
LEE D. MILLER 5
genera (only pure objective synonyms) has been attempted, nor have
the several new genera which certainly exist been named. This is
work for careful species-by-species revisions. It is certain that many
of the nomenclatorially valid genera are not biological entities; for
example, I am unable to ascertain any significant differences at the
generic level between the seven nomenclatorially valid genera included
in the satyrine tribe Melanargiini. Nevertheless, all of the nomencla-
torially valid names are carried in this revision.
For the initial work on this project the collection of Carnegie
Museum was examined for representatives of the known genera. It
soon was apparent that the material at hand was not sufficient, so
two months were spent at that Mecca of butterfly systematists, the
British Museum (Natural History), examining those genera which
were either unrepresented at Carnegie Museum or only poorly rep-
resented. Material was found and examined in these collections rep-
resenting all but four of the genera that had come to my attention by
August, 1964. Those genera that have been described since that
time are listed in what I believe to be their proper systematic posi-
tions, but none of them has been examined as critically as have the
others.
Sokal and Sneath (1963: 161-162) explain the "exemplar meth-
od" of taxonomy as follows: a taxon may be characterized by a small
number of entities within it (even a single specimen) because the
variation within taxa is considered to be less than that between taxa.
This is somewhat the method that has been employed in the present
study. The type-species of a genus is considered to be typical of it,
and the various genera are defined chiefly by their type-species in the
following pages. Occasionally the type-species is aberrant, but it is
the type-species which bears nomenclatorial "responsibility" for the
generic name, in any event. Where possible, other species of the
genera were examined, though not in as great detail as the type-
species.
The systematic revision is based entirely upon external morpho-
logical characteristics with many attributes being considered. Wing
venation drawings were made from specimens either with an ocular
grid under a dissecting microscope or with a projection device for
larger specimens. Antennae, palpi, fore-, mid- and hindlegs were re-
moved from the specimen, bleached in clorox and mounted on perma-
MEM. AMER. ENT. SOC, 24
b THE SATYRIDAE
nent slides. Drawings of these structures were made with a camera
lucida.
The taxonomic and phylogenetic analysis of the family has been
generally based on the premises that ( 1 ) entities which more nearly
resemble one another in many characteristics are most closely re-
lated, ( 2 ) structures which are reduced and fused represent advanced
conditions (Fox, 1956: 24; Fox and Fox, 1964: 71-72; Brown,
1965) and (3) those organisms which bear advanced characteristics
(not just a single advanced character, but several of them) are prob-
ably themselves advanced (for example, see Mayr, 1965). This is
in accord with the current paleontological evidence drawn from ex-
isting fossil records.
The only taxonomically valuable structures not utilized in this
study were the male and female genitalia. Examination of several
regional works (for example, Hayward, 1953; Forster, 1964) shows
little reliance can be placed on genitalic structures for classification
above the generic level, although these structures are certainly of great
importance in specific, and sometimes generic, determination.
Authorship of higher taxa has caused some problems. According
to Art. 36 of the International Code of Zoological Nomenclature
(1961), the proposition of the family Satyridae by Boisduval (1836,
as "Satyrides") implicitly creates also the subfamily Satyrinae and
the tribe Satyrini. It is this rule which has been followed through-
out the present paper for the assignment of authorship to the higher
taxa.
The zoogeographic analysis has been carried on in light of the
principles of vertebrate zoogeography, which is firmly based on and
supported by fossil evidence. For a summary of such evidence see
Darlington (1957). There are too few butterfly fossils to support or
refute any zoogeographic statement. Heavy reliance has been placed
on the presence in one area or another of relict, annectant forms:
such species may occur nearer the place of origin of the taxon than
do others. Although fossil evidence shows that such groups as horses
and rhinoceri evolved more extensively in the Nearctic than in the
Palearctic, yet are now absent from the former, many other groups
have evolved more or less in the places of their greatest present-day
density and diversity. In the absence of fossil evidence to the con-
trary, the latter situation is considered to be likely in the satyrid but-
terflies.
LEE D. MILLER
Comparative Morphology
The characters utilized in this study are all external and mor-
phological. Their distribution and evolutionary patterns are given
in the following pages.
Antennae. — Four measurements were made on the antennae:
the total length; the length of the club, where the club is a discreet
entity; the width of the shaft and the width of the club at its thickest
point. The antennae of Satyridae demonstrate the tricarinate con-
dition typical of the Nymphaloidea (Fig. 249), although the ridges
may not be apparent throughout the entire length of the antennae.
The antennae are scaled, but only on the basal few segments in the
Haeterinae. In general, the antennal club is not greatly thickened
nor flattened at the tip in the primitive satyrids (Figs. 114, 122, 128
which are some of the best developed ones), thereby following the
general primitive condition in butterflies. Many of the more advanced
Satyrinae. conversely, have the antennal club strongly developed, flat-
tened at the tips and occasionally exhibiting bizarre configurations in
some Hypocystini and Pronophilini (Figs. 163, 166, 169, 291, for
example ) . The antennal clubs of other Satyrinae are shown in Figs.
142, 151, 191, 213, 220, 235, 249, 262, 297.
Head. — Three measurements were made on the head region:
least intraocular distance; height of the eye and width of the eye.
The eyes of all satyrids are entire. Little significance could be gleaned
from these measurements, but the eyes of the Satyrinae appear to be
more ovoid than those of other subfamilies.
Labial Palpi. — In common with other butterflies the labial palpi
of the Satyridae are three-segmented. The basal segment is so firmly
attached to the head that it was often broken in dissection, hence is
little used in the present study. Five measurements were made on
the palpi: the length of the first segment, when it was not broken; the
length of the second segment; the length of the third segment; the
maximum width of the second segment and the maximum length of
the hairs of the second segment. The relationship between the lengths
of the second and third segments of the palpi varies, often without
regard to phyletic groups. For example, most of the Satyrini display
an extremely short third palpal segment, about one-fifth as long as
the second, whereas the Pronophilini. which are closely related to the
Satyrini and may have been derived from a fairly recent common an-
MEM. AMER. ENT. SOC, 24
8 THE SATYRIDAE
cestor, show the longest third palpal segment of any satyrid tribe —
in some instances over half as long as the second segment. The sec-
ond segment is generally between five and eight times as long as it is
wide, again with no apparent phylogenetic correlation. The relative
lengths of the hairs of the second segment of the palpi show some in-
teresting trends. In general the hairs are longer in the Satyrinae than
in the other subfamilies, but this tendency is modified in those groups
which have both tropical and temperate or alpine members. In such
instances the relative length of the hairs increases as the altitude or
latitude increases. Examples of the labial palpi of the Satyridae are
shown in many figures in the text, such as Figs. 3, 10, 22, 30, 35, 50,
82, 93, 100, 129, 135, 143, 172, 198, 214, 225, 236, 250, 263,
298, 311.
Thorax. — Only one thoracic measurement was made: the length
of the dorsal thoracic elements. This measurement, unfortunately, is
generally dependent only upon the absolute size of the insect.
Forelegs. — The forelegs of both sexes are more or less reduced
in all satyrids, as in other members of the Nymphaloidea. The fore-
coxae are moveable, as determined by examination of many (though
not all ) genera. The forecoxae are not reduced to the extent that the
more distal elements of the forelegs are. The reduction of the pro-
thoracic femur, tibia and tarsus has long been the criterion for re-
garding the Nymphaloidea as the most advanced of the lepidopterous
insects, a concept recently challenged by Ehrlich (1958).
Male forelegs. — Four measurements were made on the male
forelegs: the length of the tarsus; the length of the first tarsal sub-
segment; the length of the tibia and the length of the femur. The
miniaturization of the male forelegs generally follows phylogenetic
lines, being least miniaturized (and reduced; for a discussion of the
terms "miniaturized" and "reduced" see Fox, 1967) in the Haeter-
inae, Brassolinae, Biinae, Elymniinae and Eritinae and most in the
Ragadiinae and Satyrinae (compare the graphs in Figs. 1, 20, 170,
309, for example). In the first five subfamilies the femur, tibia and
tarsus usually are all relatively well developed (an exception is Cae-
rois, Fig. 27, where the tibia and the tarsus are much miniaturized).
The foretarsus in these primitive subfamilies is typically monomerous,
but some genera may have up to four freely articulating subsegments.
Examples of the forelegs of typical primitive satyrids are Figs. 4, 11,
LEE D. MILLER V
23, 51, etc. In the Ragadiinae and Satyrinae the forelegs are much
more miniaturized and reduced than in the other five subfamilies, and
occasionally there is an apparent total loss of the tarsus (Ypthimini,
Fig. 177 ). The tibia and femur are also greatly miniaturized, but the
coxa does not seem affected by this general trend. An anomalous
situation exists in the Satyrini and Erebiini, however, where a few
genera display great miniaturization of the total foreleg but still re-
tain up to four distinct, though tiny, tarsal subsegments. The male
foreleg of Ragadia is shown in Fig. 136, and those of representative
Satyrinae are shown in Figs. 144, 173, 193, 199,215,226,237,251,
266, 299, 312, etc.
Female forelegs. — Five measurements were made on the female
forelegs: the length of the tarsus; the length of the first tarsal sub-
segment; the length of the fifth tarsal subsegment, if any was present;
the length of the tibia and the length of the femur. The female fore-
legs potentially show more precise inter- and intrasubfamilial rela-
tionships than do those of the males. The forelegs of the Haeterinae,
Brassolinae, Biinae and Elymniinae are quite well developed (Figs. 1,
20, 41, 48, 88, 89, 98), always have pentamerous tarsi and usually
have posttarsal pulvilli present, except a few of the more advanced
Elymniinae. In these families the tarsus is not, or only weakly,
clubbed (for example, Figs. 5, 15, 24, 52, 112). In only two genera,
Paralethe and Antirrhea (Fig. 24) are there posttarsal claws, and
always there is but a single claw. This represents a more advanced
condition generally than that shown in the Ithomiidae (Fox, 1956:
13). The female forelegs of the Eritinae (Figs. 125, 131) and the
Ragadiinae (Fig. 137) are miniaturized beyond the point shown in
the preceding subfamilies, but they retain the full complement of
tarsal subsegments; the tarsus itself is clubbed and not so freely ar-
ticulated as in the primitive subfamilies. In the Satyrinae the fore-
legs range from miniaturized, but with five tarsal subsegments, to
highly reduced, as much so or more than in the males. Hypocystini,
Ypthimini (interestingly where the greatest reduction takes place in
the male foreleg), Euptychiini, most Pronophilini and some members
of several other tribes have pentamerous tarsi (Figs. 145, 174, 194,
200, 267, for example), but the tarsus is always clubbed. The fore-
legs of female Satyrini, Melanargiini, Dirini, Coenonymphini and
Maniolini are always reduced and apparently lack one or more tarsal
MEM. AMER. ENT. SOC, 24
10 THE SATYRIDAE
subsegments (Figs. 216, 227, 238, 252, 300, 313, for example).
The miniaturization of the forelegs of Satyrinae through the phyletic
lines may be seen by examination of Figs. 140, 170, 211, 295, 309,
etc.
Walking Legs. — Six measurements were made on both the
mesothoracic and metathoracic legs: the length of the tarsal claws;
the length of the tarsus, exclusive of the claws; the length of the first
tarsal subsegment; the length of the tibia; the length of the tibial spurs,
if any, and the length of the femur. The more important taxonomic
characters are on the mesothoracic legs. The lack of tibial spurs
characterizes the Haeterinae, Biini and Elymniini (Figs. 6, 33, 96),
and these spurs have been lost sporadically in such other groups as
the Ypthimini (Fig. 179) and Coenonymphini (Fig. 221). The
posttarsal claws are bifid in the Melanitini (Fig. 39) and in a few
other genera (Samanta and Manataria, Fig. 47). In all other Saty-
ridae the claws are simple. The relative lengths of the midtibia and
the first midtarsal subsegment are important taxonomically, particu-
larly within the Satyrinae. Usually the midtibia is between one and
a half and two times as long as the first midtarsal subsegment, but
the midtibia is barely longer than the first tarsal subsegment in the
Satyrini (Fig. 301) and in some Dirini (Fig. 259), whereas the mid-
tibia is always more than twice as long as the basal midtarsal sub-
segment in the Coenonymphini (Fig. 217). All members of the
Satyrus series of the Satyrini display a heavy spine at the dorsal distal
end of the midtibia (Fig. 301), a characteristic weakly shown by
some Dirini (as Torynesis, Fig. 259), thereby indicating a close re-
lationship between these two tribes. Dorsal spinosity of the midtibia
is shown by some members of most groups and is characteristic of
others: Haeterinae, Zetherini, Pronophilini, Dirini, Satyrini and Me-
lanargiini.
Examination of the charts of relative lengths of legs shows that
the hindlegs are occasionally significantly longer (Satyrini, Fig. 295,
for example) or shorter (the Pronophila series of the Pronophilini,
Fig. 260, for example) than the midlegs, but generally these legs are
subequal.
Wings. — The wings have long been the most-used basis for the
classification of butterflies, both from the aspect of venation and the
distribution of androconial patches. Undoubtedly too much empha-
LEE D. MILLER 1 I
sis has been placed in the past on wing venation and androconial dis-
tribution alone, but it is illogical to ignore the structure of the wings
because of previous preoccupation.
Forewings: The following four measurements were made to de-
limit the shape of the forewing: the length from the base to the end
of vein Rr,; the length from the base to the end of M2; the length from
the base to the end of 2A and the distance between the end of R-, and
the end of 2 A. The cell was delimited by two measurements: from
the base to the origins of Mi and M3. Five measurements were made
to determine the relative positions of the origins of the veins: the dis-
tance from the origin of Rs to the origin of Mi; from the origin of
Mi to the origin of M2; from the origin of M-> to the origin of M3;
from the origin of Ms to the origin of Cui and from the origin of Cui
to the origin of Cul>. Inasmuch as the inflation of the forewing veins
has long been considered a key characteristic of the Satyridae, the
following three measurements were made: the thickest part of Sc; the
thickest part of the cubital stem and the thickest part of 2A. In gen-
eral the longest forewing cell is displayed by the most primitive sub-
families, the Haeterinae (Figs. 2, 7, 8), Brassolinae (Figs. 9, 13. 14,
17, 19) and Biinae (Figs. 21, 26, 28, 29, 34, 40), where the cell is
frequently more than three-fifths the length of the wing. In most
other groups the cell is about half as long as the wing, but the Elym-
niini are characterized by an extremely short forewing cell (Figs. 90-
92), often less than a third the length of the wing. There appear to
be no broad phylogenetic implications shown by the relative place-
ment of the veins along the cell, though several relationships are shown
within subfamilies, particularly the proximity of the origins of M3
and Cui in the Elymniini (Figs. 90-92). The classic definition of
the Satyridae includes a statement about the inflation of the stems of
the three basic forewing veins. This is simply not diagnostic: the
inflation of the veins of Melanargia (Fig. 310), for example, is con-
siderably less than that of the true nymphalid genus Mestra. Within
the Satyridae particular inflation patterns are characteristic: the My-
calesini are characterized by the inflation of all three stems, that of
2A being more or less quadrate (Figs. 99, 104-106, 109, 113, 115,
119); all stems are inflated in the Hypocystini (Figs. 141, 147-150,
156, 161, 162, 165, 167, 168) and Coenonymphini (Figs. 212, 219),
but the inflation of 2A is rounded, not quadrate. In contrast, most
MEM. AMER. ENT. SOC, 24
12 THE SATYRIDAE
Biinae, Ragadiinae, Eritinae and Elymniinae (except the before-
mentioned Mycalesini ) have relatively uninflated cubital stems and
2A (Figs. 21, 29, 34, 49, 81, 90, 127, 134, for example). The pres-
ence of a vestige of forewing vein 3A is characteristic of the Haeteri-
nae (Figs. 2, 7, 8) and seems to connect them with the Ithomiidae.
Hindwings. — Five measurements were made to delineate the
shape of the hindwing and account for the majority of "tails": the
length from the base to the end of vein Rs; the length from the base
to the end of M :j ; the length from the base to the end of Cu 2 ; the
length from the base to the end of 2A and the distance from the end
of Rs to the end of 2 A. The cell was delimited by two measure-
ments: from the base to the origins of Mi and Ms. Five measure-
ments showed the relative placement of the veins along the cell: from
the origin of Rs to the origin of Mi; from the origin of Mi to the
origin of M 2 ; from the origin of Mi> to the origin of Ms; from the
origin of Ma to the origin of Cui and from the origin of Cui to
the origin of C\x-i. In all Satyrinae except Dirini (Figs. 248, 255,
256) the hindwing cell averaged greater than half the length of the
wing measured to the end of M :i (for example, Figs. 141. 171, 197,
212, 224, 234, 261, 296, 310), whereas in the other subfamilies the
hindwing cell is half or less the length of the wing, again particularly
short in the Elymniini (Figs. 90-92). The actual shape of the cell,
which is not reflected in the raw measurements, has some phyletic
significance. The cell is distally rounded in the Haeterinae (Figs. 2,
7, 8), perhaps representing a more generalized condition. In the
Biinae a gradation may be seen between a rounded cell and one pro-
duced at the origins of M 3 and Cui (Figs. 21, 26, 28, 29, 34, 40).
The Elymniinae and the "Ypthimini complex" of the Satyrinae, as well
as the Eritinae, have the cell produced at the origin of Ms (Figs. 49,
90, 99, 127, 171, 197, for example), whereas the Hypocysta series
of the Hypocystini and the "Satyrini complex" of the Satyrinae, in
general, display a migration of crossvein mrm,! outward at M2 and a
corresponding blunting of the cell (Figs. 141, 261, 296, 304, for ex-
ample).
The Ragadiinae (Figs. 134, 139) do not fit another part of the
classic definition of the Satyridae — the closure of the cells by tubu-
lar veins. In males of this subfamily the hindwing cell is closed by
vestigial, non-tubular veins, apparently only for the support of an
LEE D. MILLER 13
androconial patch; some females have the cell completely open.
Because of the migration of crossvein rao-m3, M2 arises nearer to
the origin of Ms than to the origin of Mi in some satyrine tribes; in
other groups M2 arises somewhat to much nearer Mi. Connate veins
are generally considered an advanced characteristic of the Lepidop-
tera, but in the Satyridae the opposite may be true. Hindwing veins
Ms and Cui are frequently connate in the more primitive subfamilies,
particularly the Haeterinae and the Elymniinae (for example, Figs.
2, 60, 90), but these veins are well separated in the Satyrinae (for
example, Figs. 261, 296), which are more advanced from a number
of other viewpoints, as detailed in the preceding pages. This situa-
tion is emphasized chiefly as an illustration of the independence of
characters and the danger of reliance on too few attributes, as has
been suggested by some authors (for example, Warren, 1947). Hind-
wing vein Ri of the hindwing characteristically is basally free in the
Brassolinae (Figs. 9, 13, 14, 17, 19) and Elymniini (Figs. 90-92).
Androconial Patches. — As noted previously the distribution of
androconial patches has been used extensively in the establishment of
genera (particularly by Moore, 1890-1893) — probably to too great
an extent. A few general statements may be made concerning the
distribution of these patches. There are several sites of the produc-
tion of androconia on the forewing, the most common of which is the
discal area just outside the cell from the inner margin to the origin
of Ms. This patch may be highly modified to include all or any part
of the area and generally consists of a patch of mealy scales. Hair
patches are less frequent on the forewing and are usually restricted
to the area below the cubital stem on the upper surface or along 2A
on the under surface, particularly in the Biinae and Mycalesini; re-
sultant distortion of the venation may occur in these groups. Several
areas generate androconial patches on the hindwing. Mealy patches
are generally found near the end of the cell in the area of M3 (for
example, Fig. 54). Hair patches are found in three general areas:
above the cell near the origins of Rs and Mi (for example, Fig. 73),
within the cell proper (particularly characteristic of the Elymniini,
Figs. 90-92, and Ragadiinae, Figs. 134, 139) and along the anal
veins (for example, Fig. 121). Since androconia appear sporadi-
cally within most groups they seem to have little phylogenetic signifi-
MEM. AMER. ENT. SOC, 24
14 THE SATYRIDAE
cance, though particular configurations may be characteristic of a few
tribes.
Male Genitalia. — Although these structures have been used ex-
tensively by some authors for the establishment of genera (particu-
larly by Forster. 1964), the genitalia are not considered in the pres-
ent study because of the great variability shown in such genera as
have not been split on genitalic grounds. While the genitalia are
undoubtedly indispensable for the determination of intrageneric affin-
ities, their unrestricted use as the basis for establishing genera is un-
tenable, partly because doing so creates hierarchical difficulties in
that genitalia often do not correlate with other structures, resulting
in numerous monotypic genera. Too many monobasic taxa tend to
obscure relationships within a larger group.
Systematic Revision of the Genera of the Satyridae
Before meaningful statements may be made concerning evolution,
phylogeny and zoogeography, it is necessary to analyze relationships.
No usable taxonomy presently exists for the Satyridae; the attempt in
the following pages to apportion genera to subfamilies and tribes is
designed to fill this need. This classification is based on many char-
acters in the hope that it may more nearly approximate a "natural"
classification than have some schemes in the past which were based
on one or very few characteristics.
It is the purpose of this revision to define, as closely as possible,
the subfamilies and tribes which comprise the Satyridae, as well as to
define the family itself. No single character can or should be ex-
pected to be diagnostic of an entire group: evolution modifies char-
acters indiscriminately, without regard to "sacred" taxonomic ones.
That a single character — or even several — is atypical of the group
in some genera is no reason to assign such genera to other tribes
(Simpson, 1961a: 54-63). The practice of "splitting' across natural
boundaries comes from the indiscriminate analysis of too few char-
acters.
Two other nymphaloid families are considered to be systemati-
cally and phylogenetically close to the Satyridae: the Morphidae and
the Ithomiidae.
The Morphidae, which are represented in both the Old and New
World tropics, have been regarded by many authors (e.g., Ehrlich,
1958) to be very near the Satyridae. With few exceptions the hind-
LEE D. MILLER 15
wing cell of the morphids is open between veins M- and M :i , whereas
only satyrids of the subfamily Ragadiinae have the cell closed by
other than fully functional tubular veins. The structure of the mor-
phid legs is substantially that shown by the primitive satyrids, but
larval foodplant preference will separate some morphids. All the
Indo-Australian morphids feed as larvae on monocotyledons, as do
the satyrids, but the Neotropical members feed predominantly on di-
cotyledons. There are a few nymphalid genera that utilize mono-
cotyledons, but Morpho is unique among nymphaloids in commonly
using both monocots and dicots (Ehrlich and Raven, 1965). On the
larval foodplant preference and on pattern morphology (see Schwan-
witsch, 1924) the New World morphids appear to be very close to
the condition apparent when the fundamental dichotomy of the nym-
phaloids into monocotyledon and dicotyledon feeding groups took
place.
Fox (1956: 18-19) considers the Ithomiidae to be as near or
nearer the Satyridae than to the Danaidae, with which the ithomiids
had formerly been placed. This conclusion was reached from the
evidence of various characteristics, particularly of the adults and pu-
pae. In many respects the ithomiids resemble the Haeterinae of the
Satyridae, particularly as regards the presence of a vestige of fore-
wing vein 3 A (a characteristic also shared with the danaids) in both
groups. The obviously similar tendency of the ithomiids and the
haeterines to have hyaline wings is, therefore, supported by structural
evidence. The vestige of forewing vein 3A will serve to separate the
ithomiids from the remainder of the satyrids, and no discreet andro-
conial areas are found in the haeterines, but these are usual (and
taxonomically important) in the ithomiids. All Ithomiidae feed as
larvae on the dicotyledonous family Solanaceae. The morphids ap-
pear to be even more closely related to the satyrids than the ithomiids,
but neither can be combined with the satyrids.
Family SATYRIDAE Boisduval, 1836
Satyrides Boisduval, 1836: 166.
= Hipparchiadae Kirby, 1837: 297.
= Satyridae Swainson, 1840: 86.
As mentioned in the preceding section, the Satyridae are related
MEM. AMER. ENT. SOC, 24
16 THE SATYRIDAE
to the nymphaloid families Morphidae and Ithomiidae.
No single key character will serve to separate the satyrids from
all other nymphaloids, but a general definition of the Satyridae may
be formulated as follows: a nymphaloid family having a larva with
a bifid tail (fleshy anal projections from the eleventh abdominal seg-
ment) and feeding on monocotyledonous plants and generally having
adults with the cells of the fore- and hindwings closed by tubular
veins, forewing veins swollen at their bases and with more or less
shaggy palpi. Only the habit of feeding on monocots and the bifid
larval tail are universal in the Satyridae, and some morphids feed on
monocotyledons. Many other nymphaloids (danaids, ithomiids, ac-
raeines, etc.) have the cells of both wings closed by tubular veins,
and the satyrid subfamily Ragadiinae is defined by not having the
hindwing cell closed by tubular veins, often not closed at all. While
many of the satyrids have shaggy palpi, many of the primitive ones
have only very short hairs on the palpi. This part of the definition
universally applies only to the temperate and arctic species. The
forewing veins are usually, but not universally, swollen at their bases.
The bases of the forewing veins of such satyrid genera as Melanitis,
Lethe, Oeneis and Melanargia are less swollen than are those of such
"true" nymphalid genera as Mestra, Bolboneura and Callicore (see
Schatz andRober, 1892).
The Satyridae are nearly cosmopolitan, being found wherever but-
terflies in general are found, except on some oceanic islands. These
butterflies, like most others, find their greatest development and pro-
liferation in the tropics, not in the temperate zone as claimed by Ehr-
lich and Raven, (1965: 589).
Seven subfamilies are recognized in this family. The most primi-
tive of these are the Haeterinae, Brassolinae and Biinae; intermediate
are the Elymniinae and Eritinae; the Ragadiinae and Satyrinae are
advanced.
In the following pages the individual subfamilies are characterized
and diagnoses are provided for the tribes embodying the following
parameters: (1) condition of the eyes; (2) relative length of the an-
tennae; (3) characteristics of the antennal club; (4) relative lengths
of the second and third segments of the palpus; (5) the development
of the hairs of the palpus; (6) the development of the male foreleg;
(7) the development of the female foreleg; (8) the relative lengths
LEE D. MILLER
17
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18 THE SATYRIDAE
of the mid- and hindlegs, if significant; (9) the relative lengths of the
midtibia and the first midtarsal subsegment; (10) the presence or ab-
sence of dorsal spinosity on the midtibia; (11 ) the development of
tibial spurs, if any; (12) the presence or absence of a dorsal, distal
spine at the end of the midtibia; (13) the length and shape of the
fore wing cell; (14) the branching pattern of the forewing radial veins;
(15) the relative positions of the origins of torewing veins Rs and
Mi; (16) the relative positions of the forewing medial veins; (17)
the relative positions of forewing veins Ms, Cui and Cuy, (18) the
patterns of inflation of the forewing vein bases; (19) the length and
shape of the hindwing cell; (20) the relative lengths of hindwing
veins Sc + Ri and 3 A; (21) the relative positions of hindwing veins
Mh and Cui; (22) the relative positions of the hindwing medial veins;
and (23) characteristic and aberrant superficial patterns and the dis-
tribution of androconia.
Table 2 gives the character states of certain key characteristics
within the satyrid subfamilies. A key to these subfamilies follows.
Key to the Subfamilies of the Satyridae
1. Hindwing cell open or closed only by aberrantly placed veins (Figs. 134,
1 39) Ragadiinae
Hindwing cell closed by tubular veins (Fig. 2, tor example) 2
2. Forewing with a vestige of vein 3A at base (Figs. 2, 7, 8) Haeterinae
Forewing vein 3 A fused with 2A throughout its entire length (Fig. 21, for
example) 3
3. Hindwing vein R x free basally from Sc + R x (Figs. 9, 90, for example) 4
Hindwing vein R x completely fused with Sc (Fig. 21, for example) 5
4. Forewing cell greater than half length of wing; Neotropical species
Brassolinae
Forewing cell much less than half length of wing; Paleotropical species ....
Elymniinae (Elymniini)
5. Posttarsal claws bifid (Figs. 39. 47) 6
Posttarsal claws simple 7
6. Forewing cell more than half length of wing Biinae (Melanitini)
Forewing cell less than half length of wing
Elymniinae {Samanta) and Manataria
7 '. Female foretarsus with fewer than five subsegments many Satyrinae
Female foretarsus with five subsegments 8
8. Female foretarsus strongly clubbed distad (Fig. 174, for example)
many Satyrinae
Female foretarsus not so clubbed (Figs. 24, 52. 124, for example) 9
LEE D. MILLER 19
9. Forewing cell half length of wing or less most Eiymniinae
Forewing cell greater than half length of wing 10
10. Forewing cell produced at the origin of M 3 (Figs. 21, 29, for example);
New World species New World Biinae
Forewing cell produced at the origin of M 1 (Figs. 121, 127): Indo-Aus-
tralian species Eritinae
Subfamily HAETERINAE Herrich-Schaffer, 1864
Hetaerina [sic] Herrich-Schaffer, 1864: 124.
= Pierellinae Clark, 1947: 149.
The Haeterinae on several counts are the most primitive of the
Satyridae. These butterflies cannot be considered close to any other
subfamily but show more affinities (albeit slight ones) with the Biinae
than with the Brassolinae or Eiymniinae. From several aspects —
the free proximal end of forewing vein 3A, the general elongate wing
form, the seta-like modification of the wing scales and the preference
for a deep forest habitat — this subfamily seems to form a connecting
link with the Ithomiidae (Fox, 1956: 18-19), but, again, this is a
very remote relationship, if not a completely spurious one arising from
ecological adaptations.
In itself the free proximal vestige of forewing vein 3A serves to
distinguish the haeterines from all other satyrids. This characteristic
is otherwise shown in the Nymphaloidea by the Ithomiidae, Danaidae,
some Morphidae (Taenaris) and such nymphalid genera as Colaenis,
Kallima, Apatwina and the very primitive Callinaga. The general
shape of the wings (Figs. 2, 7, 8) is characteristic, as is the rounded
hindwing cell. The combination of a dorsally spinose midtibia and
the lack of tibial spurs (Fig. 6) is shown by no other satyrid. The
lack of distal scaling on the antennae is also peculiar to the present
subfamily.
All members of the Haeterinae are restricted to the American
tropics and are found only in the dense forests.
A single tribe, the Haeterini, includes all members of the sub-
family.
The most comprehensive revision of the Haeterinae presently
available is that of Brown (1942).
MEM. AMER. ENT. SOC, 24
20
THE SATYRIDAE
Tribe Haeterini Herrich-Schaffer, 1864
(Figures 1-8)
Hetaerina [sic] Herrich-Schaffer, 1864: 124.
= Pierellinae Clark, 1947: 149.
The diagnostic features of this tribe are those outlined for the
subfamily above. The Haeterini are characterized as follows:
The eyes are naked. The antennae vary little in length from
slightly over one-third (Pierella) to just over two-fifths (Haetera)
the length of the forewing costa. The antennal club is weakly de-
veloped, generally less than twice the thickness of the shaft and oc-
cupying the distal one-fourth to one-fifth of the antenna. The anten-
nae of the haeterines are scaled on only the proximal few segments:
the antennae of all other satyrids are fully scaled. The third segment
of the palpus is usually less than one-fifth the length of the second,
although the third segment is between one-third and one-fourth the
length of the second in Haetera and Pseudohaetera. The hairs on
the second segment of the palpus are generally about as long as the
segment is wide (the hairs are twice as long as the segment is wide
in D nice do).
0.4 x ML, FL
Haetera
Cithaerias
Pierella
Fig. 1. Haeterinae: Haeterini. Relative lengths of the femur + tibia +
tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML
value is unity. The top bar for each genus represents the measurements ob-
tained from males, the bottom bar those from females.
The male foreleg is well developed for a nymphaloid, the tibia is
longer than the femur, although both are about equal in Pierella, and
there is a single, unspined tarsal subsegment in all genera. The female
foreleg is also well developed with a pentamerous, unclubbed tarsus
LEE D. MILLER
21
Figs. 2-8. Haeterinae: Haeterini. 2. Haetera piera (Linne), 6 venation.
3. H. piera, palpus (in this and all succeeding illustrations of palpi only the
distal two segments are figured). 4. H. piera, S foreleg (in this and all suc-
ceeding illustrations of the $ forelegs, unless otherwise indicated, the femur,
tibia and tarsus are figured. 5. H. piera, 2 foretarsus. 6. H. piera, midleg
(in this and all succeeding illustrations of the midlegs only the femur, tibia
and tarsus are figured). 7. Pierella nereis (Drury), $ venation. 8. Cithae-
rias pireta (Cramer), 6 venation.
armed with spines on the second, third and fourth subsegments. The
mid- and hindlegs are of about equal length. The midtibia is one
and one-half to two times as long as the first midtarsal subsegment, is
spiny dorsad and lacks tibial spurs or a spine at the dorsal, distal end.
The relative development of the legs is shown in Fig. 1 .
The forewing cell varies from just greater than half to more than
three-fifths the length of the forewing costa and is rounded distad in
all genera. The forewing radial veins arise in three branches from
the cell. Forewing veins Rs and Mi are connate or approximate at
MEM. AMER. ENT. SOC, 24
22 THE SATYRIDAE
their origins, vein M2 arises midway between Mi and M3 and vein
Cui arises midway between M 3 and C112, or slightly nearer the former.
The forewing vein Sc is inflated in all genera, and the other veins may
be: both the cubital stem and 2A in Pierella, only 2A in Haetera.
The presence of a proximal vestige of forewing vein 3A is diagnostic
in the Haeterinae.
The hindwing cell is also distally rounded and is from two-fifths
{Pierella) to three-fifths (Cithaerias) as long as the wing, as measured
to the origin and end of M3, respectively. Hindwing veins Sc + Ri
and 3A are of about the same length. The origins of hindwing veins
M3 and Cui may be widely separated ( Cithaerias, Fig. 8 ) to connate
on a common stalk (Pierella, Fig. 7 ). Vein M L » arises about midway
between Mi and M3, or slightly nearer the latter.
The pattern is distinctive (see Schwanwitsch, 1925 ) : the scales of
most species are reduced to narrow, hairlike vestiges, resulting in a
hyaline or translucent appearance in most species. Relatively un-
modified scales are displayed by some species, particularly of the
genera Pierella and Haetera, with resultant opaque wings.
Genera Included in the Haeterini
Callitaera Butler, 1868b: 101. Type-species: "Papilio aurora Felder" (= Ci-
thaerias phantoma Fassl, fide Clifton, in litt.), designated by Brown,
1942: 311.
Cithaerias Hiibner, [1819] (1816-1826): 53. Type-species: "Papilio andro-
meda Fabricius" ( = Papilio pireta Cramer, according to Brown, 1942),
designated by Scudder, 1875a: 143. Papilio menander Drury, often
regarded as the type-species, was shown by d'Almeida (1951: 496) to
be identical with Papilio pireta Cramer.
Dulcedo d'Almeida, 1951: 501. Type-species: Haetera polita Hewitson, by
original designation.
Haetera Fabricius, 1807: 284. Type-species: Papilio piera Linne, designated
by Butler, 1868a: 195.
= Oreas Hiibner, [1807] (1806-1838): pi. [82]. Type-species: Pa-
pilio piera Linne, by monotypy. Oreas is sunk to Haetera Fabri-
cius in accordance with Opinion 137 (1942) of the International
Commission of Zoological Nomenclature.
Pierella Herrich-Schaffer, 1865: 65. Type-species: Papilio nereis Drury, desig-
nated by Butler, 1868a: 195.
Pseudohaetera Brown, 1942: 330. Type-species: Haetera hypaesia Hewitson,
by original designation.
LEE D. MILLER 23
Subfamily BRASSOLINAE Boisduval, 1836
Brassolides Boisduval, 1836: 166.
The brassolines have long been considered a separate family
somewhat intermediate between the satyrids and the morphids, but
Ehrlich (1958: 351-352) placed the brassolines with the morphids.
There is little except the large size of the butterflies to support this
combination: the larvae of most Neotropical morphids feed on dicots,
whereas the brassolines feed exclusively on monocots; brassoline lar-
vae have bifid tails, but the tails are greatly reduced in the morphids.
These two characters intimately associate the brassolines with the
other satyrids. The cells of both wings are closed in the brassolines,
as in the satyrids generally, whereas the hindwing cell is open in most
of the Morphidae. Within the Satyridae the brassolines are allied to
the New World Biinae, particularly through such genera as Narope.
There seems little justification, in view of the above facts, for retain-
ing the brassolines as a family separate from the Satyridae.
The brassolines may be distinguished from all other New World
Satyridae by the basal separation of hindwing veins Sc and Ri (Figs.
9, 13, 14, 17, 19). This characteristic is also shown by the Old
World Elymniini (Elymniinae), but the extremely short cell and vir-
tually connate veins M3 and Cui of the forewing immediately disting-
uishes the elymniines. The Haeterinae show an approximate condi-
tion of hindwing veins Sc + Ri and Rs, but there is no basal separation
of Ri. The brassolines lack the vestige of forewing vein 3 A which
characterizes the Haeterinae.
All brassolines are Neotropical, none being found further north
than southern Mexico. Some species of Caligo are banana pests in
Central America, and around the banana plantations these insects
often may be seen flying just at dusk.
Clark ( 1947, 1948 ) recognized two tribes as subfamilies of "Bras-
solidae", but no good evidence is found for splitting the brassolines
into more than a single tribe, the Brassolini. Within the Brassolini
groupings are possible, but not tribal ones.
Tribe Brassolini Boisduval, 1836
(Figures 9-19)
Brassolides Boisduval, 1836: 166.
= Caliginae Clark. 1947: 149.
MEM. AMER. ENT. SOC, 24
24
THE SATYRIDAE
Figs. 9-19. Brassolinae: Brassolini. 9. Brassolis sophorae (Linne), £
venation. 10. B. sophorae, palpus. 11. B. sophorae, £ foreleg. 12. B. so-
phorae, midleg. 13. Opsiphanes cassiae (Linne), £ venation (after Schatz
and Rober, 1892). 14. Caligo atreus Kollar, £ venation (after Schatz and
Rober, 1892). 15. C. eurylochus (Cramer), 9 foretarsus. 16. C. eiirylochus,
midtarsus and distal portion of midtibia. 17. Dasyophthalma rusiiui (Godart),
£ venation. 18. D. rusina, 9 foretarsus. 19. Narope cyllastros Doubleday
and Hewitson, £ venation.
The characteristics of the subfamily are those of the tribe. Four
groups may be noted within the tribe, but they are only loose as-
semblages and tend to grade into one another. The diagnostic char-
acters are summarized in Table 3.
LEE D. MILLER
25
The Brassolis-series, sensu stricto, is composed of robust species
with naked eyes, moderately well developed or absent tibial spurs and
a basically brown upper surface with a transverse band of orange,
yellow or red across the forewing.
The Caligo-senes includes large, broad-winged, but not robust-
bodied butterflies with naked eyes, well developed tibial spurs and a
basically iridescent purple upper surface with faint longitudinal light
markings.
The Dasyophthalma-series is comprised of a single genus of broad-
winged species with hairy eyes, moderately well developed tibial spurs
and a basically brown upper surface, with or without iridescence, and
with longitudinal lighter markings.
The Narope-sevies is also comprised of a single genus of medium-
sized butterflies with acute forewing costal margins, naked eyes, mod-
erately well developed tibial spurs and a brown upper surface with
little or no maculation.
Table 3
The character states of four characters ir
i the genera
of the Brassolinae
Brassolini.
Length of fore-
wing cell/
Antennal
Tibial
length of
Genus
Eyes
club
spurs
forewing costa
Brassolis-series, s. str.
Brassolis
Naked
Pronounced
Present, ex.
3 5-2/3
B. sophorae
Catoblepia
Naked
Gradual
Present
3 5-2/3
Dynastor
Naked
Gradual
Present
1/2-3/5
Opoptera
Naked
Gradual
Present
3/5-2/3
Opsiphanes
Naked
Gradual
Present
1/2-3 5
Penetes
Naked
Gradual
Present
3/5-2/3
Selenophanes
Naked
Gradual
Present
3/5-2/3
Caligo-senes
Caligo
Naked
Gradual
Present
3 5-2/3
Eryphanis
Naked
Gradual
Present
3/5-2/3
Dasyophthalma-series
Dasyophthalma
Hairy
Gradual
Present
3 5-2/3
Narope-series
Narope
Naked
Gradual
Present
just >l/2
MEM. AMER. ENT. SOC, 24
26 THE SATYRIDAE
The Brassolini are characterized as follows. — The eyes are naked
in all genera but Dasyophthalma in which they are densely hairy.
The antennae are short, one-third to two-fifths the length of the fore-
wing costa, and the antennal club is weakly developed in most genera
(just over twice the width of the shaft in Brassolis). The third seg-
ment of the palpus is generally one-fourth to one-sixth the length of
the second, although it is far shorter in such rather advanced genera
as Brassolis and Opsiphanes and somewhat longer in the more primi-
tive Narope. The hairs of the second segment of the palpus are gen-
erally shorter than the segment is wide.
The male foreleg is well-developed with a mono- to trimerous tar-
sus; the femur is longer than the tibia in all genera. The female fore-
leg is likewise well developed with a pentamerous, unclubbed tarsus
bearing spines on at least the third and fourth (often the first and
second, too) subsegments. The midtibia varies from just less than
twice {Caligo and Eryphanis) to well over twice ( Brassolis-series )
as long as the proximal midtarsal subsegment. The midtibia is spiny
dorsad with well to moderately well developed tibial spurs (except in
Brassolis sophorae, where the tibial spurs are totally wanting, Fig.
12); there is no spine at the dorsal, distal end of the midtibia.
The forewing cell is produced at the origin of Ma, particularly in
Caligo and Eryphanis. The cell is generally three-fifths to two-thirds
the length of the forewing costa, although it is somewhat shorter in
Dynastor, Opsiphanes and Narope. The forewing radial veins arise
from the cell in three branches. Forewing veins Rs and Mi are well
separated at their origins; vein M L > usually arises nearer Mi than M3,
although M2 arises nearer M3 in Caligo and Dynastor; vein Cui arises
nearer Ma than CU2, except in Dasyophthalma. The forewing veins
are seldom inflated, though the base of Sc may be in some genera.
The hindwing cell is produced marginad by a migration of cross-
vein mriii3 and is generally about three-fifths the length of the wing,
measured to the origin and the end of Ma, respectively. Hindwing
veins Sc and Ri are separate proximally in this tribe, as in the Elym-
niini (Elymniinae); Sc + Ri is longer than 3A. At their origins veins
Ms and Cui are well separated, and M 3 arises nearer Ma than Mi.
The upper surface patterns have been described in the analysis of
the generic series. On the under surface the wings are cryptically
patterned, light brown with darker striations, much as in the Antir-
LEE D. MILLER 27
rhini. In the brassolines, however, there are often large "eye-spots",
particularly on the hindwing, which have been credited with being
protective markings (the wings are rapidly opened when the butter-
fly is threatened by a predator, giving the impression of large eyes
blinking). The development of these eye-spots gives the common
name of the Cali go-series, the "owl butterflies".
Genera Included in the Brassolini
The Brassolis-series, sensu strict o
Brassolis Fabricius, 1807: 282. Type-species: Papilio sophorae Linne, desig-
nated by Westwood, 1851, in Doubleday, Westwood and Hewitson,
1846-1852: 341.
Catoblepia Stichel, 1902: 488, 491. Type-species: Brassolis amphiroe Hiibner,
by original designation.
Dynastor Westwood and Hewitson, 1849, in Doubleday, Westwood and Hewit-
son, 1846-1852: pi. 58. Type-species: Dynastor napoleon Westwood
and Hewitson, by monotypy.
Opoptera Aurivillius, 1882: 75. Type-species: Brassolis syme Hiibner, by origi-
nal designation.
Opsiphanes Westwood and Hewitson, 1849, in Doubleday, Westwood and Hew-
itson, 1846-1852: pi. 57. Type-species: Opsiphanes sallei Westwood
and Hewitson, designated by Scudder, 1875a: 233.
Penetes Westwood and Hewitson, 1849, in Doubleday, Westwood and Hewit-
son, 1846-1852: pi. 58. Type-species: Penetes pamphanis Westwood
and Hewitson, by monotypy.
Selenophanes Staudinger, 1888: 212. Type-species: Papilio cassiope Cramer,
by original designation.
The Caligo-sehes
Caligo Hiibner. [1819] (1816-1826): 51. Type-species: Caligo eurylochus
Hiibner, designated by Scudder, 1875a: 129.
Eryphanis Boisduval, 1870: 57. Type-species: Papilio automedon Cramer, by
monotypy.
Dasyophthalma-series
Dasyophthalma Westwood, 1851, in Doubleday, Westwood and Hewitson,
1846-1852: 343. Type-species: Brassolis rusina Godart, designated by
Scudder, 1875a: 155.
MEM. AMER. ENT. SOC, 24
28 THE SATYRIDAE
/Varo/^-series
Narope Westwood and Hewitson, 1 849, in Doubleday, Westwood and Hewit-
son, 1846-1852: pi. 50. Type-species: Narope cyllastros Westwood
and Hewitson, by monotypy.
Subfamily BIINAE Herrich-Schaffer, 1864
Biina Herrich-Schaffer, 1864: 124.
The Biinae are the third of the very primitive satyrid groups and
show no especially close relationship with any other subfamily. The
Neotropical biines share some vague affinities with the haeterines and
some firmer ones with the brassolines, particularly as regards vena-
tion, whereas the Paleotropical biines grade toward the Lethini of the
next subfamily.
The biines are generally large to very large satyrids, often brightly
colored above and always cryptically patterned beneath with few and
usually poorly developed eye-spots on either wing surface. The long
forewing cell, generally three-fifths the length of the wing except in
some melanitines (Fig. 34), is characteristic, and the well separated
hindwing veins M3 and Cui serve to distinguish these butterflies from
most elymniines. The wings are never translucent or transparent, as
in the haeterines.
Table 4
Some of the diagnostic characters of the tribes of the Biinae.
Character Antirrhini Biini Melanitini
Tarsal claws Simple Simple Bifid
Tibial spurs Present, but Absent Present
poorly devel-
oped
Length of 3rd seg. of
palpus relative to
length of second <l/4 >l/2 1/4
Forewing radial veins 3 branches 1 branch, 3 branches
aberrant
Inflation of forewing veins .. Only Sc, if any; .. Sc, Cu, 2A Only Sc, slight
or else v. slight greatly inflated
LEE D. MILLER 29
All biines are tropical. The Antirrhini and Biini are found ex-
clusively in the Neotropics and are forest dwellers, whereas the Mel-
anitini are Paleotropical (from West Africa to Australia) and are
found in less dense situations, though many of these species are cre-
puscular.
The Biinae are split, somewhat reluctantly, into three tribes, but
Bia is far too aberrant to be referred to either of the other two. The
diagnostic features of the three tribes are given in Table 4, and a key
to these tribes is given below.
Key to the Tribes of the Biinae
1. Tarsal claws bifid (Fig. 39); Paleotropical species Melanitini
Tarsal claws simple; Neotropical species 2
2. Tibial spurs present, though poorly developed (Fig. 25); third segment of
palpus less than one-fourth length of second segment (Fig. 22)
Antirrhini
Tibial spurs absent (Fig. 33); third segment of palpus greater than half
length of second segment (Fig. 30) Biini
The New World members of the Biinae are considered to be more
primitive because of ( 1 ) their greater convergence toward the Bras-
solinae and Haeterinae, (2) the greater relative size of the male fore-
leg and (3) the unclubbed female foreleg. The Melanitini are the
more advanced members of the subfamily and seem to show relation-
ships with the more advanced Elymniinae.
Tribe Antirrhini, new tribe
(Figures 20-28)
The simple tarsal claws serve to separate this New World tribe
from the Old World Melanitini. Many characteristics serve to sep-
arate the Antirrhini from the Biini, notably the presence of tibial
spurs, the more reduced third segment of the palpus, the arising of
the forewing radial veins in three branches and the relative placement
of several other veins on the cells.
The Antirrhini are Neotropical insects, found from Guatemala to
Paraguay and northern Argentina. All seem to be forest species,
seldom venturing into open areas. The tribe Antirrhini is character-
ized as follows:
The eyes are naked in all genera but Sinarista, in which they are
sparsely hairy. The antennae are about half as long as the forewing
MEM. AMER. ENT. SOC, 24
30
THE SATYRIDAE
0.4xML,
FL
— ■+
l_
x;
'///////>
ML
ANTIRRHINI
HL
BIINI
IxML
Antirrhea
Caerois
Triteleuta
Bia
Melanitis
Hipio
Cyllogenes
Parantirrhoea
Fig. 20. Biinae: Antirrhini, Biini and Melanitini. Relative lengths of the
femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light
stippling) and hindlegs (HL, heavy stippling) of selected genera. In all in-
stances the ML value is unity. The top bar for each genus represents the mea-
surements obtained from males, the bottom bar those from females.
costa, and the antennal club is weakly developed (it is completely in-
distinct in Triteleuta). When the club is distinct it occupies the dis-
tal one-third to one-quarter of the antenna and is seldom more than
twice as thick as the shaft. The third segment of the palpus is one-
fourth to one-sixth the length of the second segment, and the hairs of
the second segment are about as long as the segment is wide, although
the hairs of Sinarista are about twice as long as the width of the seg-
ment.
The male foreleg is well developed with a monomerous, unspined
tarsus, and the tibia is usually slightly longer than the femur, although
the femur is much longer in Caerois. The female foreleg is also well
developed with a pentamerous, unclubbed tarsus bearing spines on
the first four subsegments. The midtibia is more than twice as long
LEE D. MILLER 31
as the basal midtarsal subsegment, smooth dorsally and bearing weakly
developed tibial spurs. There is no spine at the dorsal, distal end of
the midtibia. The relative lengths of the legs are shown in Fig. 20.
The forewing cell is square-cut in Caerois and Sinarista, but deeply
excavate at Mi in the other genera. The cell is about three-fifths the
length of the forewing costa. The radial veins arise in three branches
from the forewing cell, and veins Rs and Mi are connate to approxi-
mate at their origins. Forewing vein M- arises midway between Mi
and M ;i , whereas Cu t usually arises about midway between M:t and
CU2 (nearer M3 in Caerois). The forewing veins are only slightly
inflated, if at all; any thickening is at the base of Sc.
The hindwing cell is highly variable, from very short and narrow
in Antirrhea to over three-fifths the length of the wing, measured to
the end of M3, and rounded in Caerois. Hindwing vein Sc + Ri is
usually longer than 3A, although the two veins are of about the same
length in Sinarista. Hindwing veins M3 and Cui are generally well
separated at their origins, but they are approximate in Antirrhea; vein
M2 arises midway between Mi and M3.
Most of the species are rather brightly colored above with few
ocelli, although some species, such as Sinarista adopt iva Weymer,
have rows of submarginal ocelli. On the under surface all species
are cryptically marked with many fine, dark striations on a tan to
brown ground color. There are androconial hair patches on the under
surface of the forewing along vein 2 A in all genera but Sinarista, and
these androconial areas frequently result in distortions of the venation.
Genera Included in the Antirrhini
Antirrhea Hiibner, [1822] (1806-1838): pi. 294. Type-species: Antirrhea ar-
chaea Hiibner, by monotypy.
= A nchiphlebia Butler, 1868b: 106. Type-species: Antirrhea archaea
Hiibner, by original designation.
Caerois Hiibner, [1819] (1816-1826): 56. Type-species: Papilio arcesilaus
Cramer (= Papilio chorinaeus Fabricius). by monotypy.
= Arpidea Duncan, 1837: 180. Type-species: Papilio chorinaeus
Fabricius, by monotypy.
Sinarista Weymer, 1909: 164. Type-species: Sinarista adoptiva Weymer, by
monotypy.
Triteleuta Strand, 1912: 44. Type-species: Antirrhea tomasia Butler, by origi-
nal designation. Proposed as a subgenus of Antirrhea Hiibner.
MEM. AMER. ENT. SOC, 24
32
THE SATYRIDAE
Figs. 21-28. Biinae: Antirrhini. 21. Antirrhea archaea Hiibner, $ vena-
tion. 22. A. archaea, palpus. 23. A. archaea, $ foreleg. 24. A. archaea, 9
foretarsus. 25. A. archaea, midleg. 26. Caerois chorinaeus (Fabricius), $
venation. 27. C. chorinaeus, $ foreleg. 28. Sinarista adoptiva Weymer, $
venation.
Tribe Biini Herrich-Schaffer, 1864
(Figures 20, 29-33)
Biina Herrich-Schaffer, 1864: 124.
Bia actoriaena (Linne), the only species included in the Biini, is
the most aberrant member of the Satyridae. It has been considered
variously a brassoline (Weymer, 1912; Clark, 1947, 1948) or a per-
fectly good satyrid (Rober, 1892; Ehrlich, 1958). Bia is definitely
LEE D. MILLER 33
a satyrid and shows relationships to other members of the Biinae,
chiefly through pattern convergence, but it is far too aberrant to be
included in either of the other two biine tribes. This species can be
distinguished from all other biines by the lack of tibial spurs, in addi-
tion to a multitude of other characteristics enumerated below in the
description of the tribe. Bia is primitive, though highly specialized,
and may be characterized as follows:
The eyes are naked. The antennae are half as long as the fore-
wing costa, and the club is poorly developed occupying the distal
quarter of the antenna and about twice as thick as the shaft. The
third segment of the palpus is very long, over half the length of the
second segment, and the hairs of the second segment are not as long
as the segment is wide.
The male foreleg is well developed, with the femur longer than the
tibia and a monomerous, unspined tarsus. The female foreleg is also
well developed with a pentamerous, unclubbed tarsus bearing spines
on the first four subsegments. The midtibia is smooth dorsally, more
than twice as long as the proximal midtarsal subsegment and armed
with neither tibial spurs nor a spine at the dorsal, distal end.
The forewing cell, measured to the origin of M3, is greater than
three-fifths the length of the forewing costa, but the cell is deeply ex-
cavate at the origins of Mi and M2, and the cell is only about half
the length of the forewing costa when it is measured to the origin of
Mi. All the forewing radial veins arise from a single branch from
the cell, and Ri anastomoses distally with Sc, resulting in a confused
venational pattern along the costa. Forewing veins Rs and Mi are
well separated at their origins. Vein M2 arises much nearer Mi than
M s , whereas Cui arises midway between M3 and Cui>. Veins Sc, the
cubital stem and 2A are all strongly inflated at their bases, the in-
flation of the latter suggesting the pattern shown in the Mycalesini of
the next subfamily.
The hindwing cell is strongly produced at the origin of Ms and
about three-fifths the length of the wing measured to the end of M3.
Hindwing vein Sc + Ri is longer than 3 A. Veins M3 and Cui are
well separated at their origins, and Ml> arises nearer Mi than M3.
The upper surface is dark brown with patches of orange and iri-
descent blue scales. On the under surface Bia is marked like other
members of the subfamily: tan with numerous, longitudinal brown
MEM. AMER. ENT. SOC, 24
34
THE SATYRIDAE
Figs. 29-33. Biinae: Biini, all Bia actoriaena (Linne).
30. palpus. 31. A foreleg. 32. 2 foretarsus. 33. midleg.
29. £ venation.
striae. The androconial areas on the hindwing are characteristic:
there is a patch of mealy scales on the upper end of the cell along
crossvein r s -nii, and a long hair tuft lies along 2A.
Genus Included in the Biini
Bia Hubner, [1819] (1816-1826): 51. Type-species: Papilio actoriaena Linne,
by monotypy.
Tribe Melanitini, new tribe
(Figures 20, 34-40)
The melanitines are the most advanced members of the Biinae, as
noted above, particularly with regard to the more reduced male fore-
leg and the slightly clubbed female foretarsus (Fig. 37). There are,
in addition to the already-mentioned affinities to the Antirrhini, some
rather suggestive ties between the Melanitini and the Lethini of the
next subfamily, especially through the genera Parantirrhoea (Mela-
nitini) and Ptychandra and Samanta (Lethini). The venation of
Pty chandra and its pattern relate rather closely (but with modifica-
tions) to those of the Melanitini, and Samanta displays the bifid claws
characteristic of the melanitines. Bifid claws are the most character-
istic feature of this tribe and serve to distinguish it from all other
satyrids except Samanta and Manataria (a Neotropical genus of un-
certain position).
LEE D. MILLER
35
All members of this tribe are Paleotropical, being found from
West Africa through the Malay Peninsula east as far as New Guinea
and northern Australia. At least some of the species are crepuscular,
and they do not appear to be so limited to the deep forest as are the
Neotropical Antirrhini. The Melanitini are characterized as follows:
The eyes are naked. The antennae are rather uniformly just over
two-fifths the length of the forewing costa, and the antennal club is
weakly developed, occupies the distal quarter of the antenna and is
thickened twice to two and a half times the thickness of the shaft.
The third segment of the palpus is about one-fourth the length of the
second, and the hairs of the second segment are much shorter than
the segment is wide.
Figs. 34-40. Biinae: Melanitini. 34. Melanitis leda (Linne), 6 venation.
35. M. leda, palpus. 36. M. leda, S foreleg. 37. M. leda, 2 foretarsus. 38.
M. leda, midleg. 39. M. leda, bifid tarsal claw. 40. Hipio constantia (Cramer),
venation of proximal portion of 6 wings.
The male foreleg is well developed with the femur and the tibia
of about the same length (the femur is longer in Cyllogenes and
Parantirrhoea) and bearing a single, usually unspined (weakly so in
Cyllogenes) tarsal subsegment. The female foreleg is also well de-
veloped with a pentamerous, unclubbed or weakly clubbed tarsus.
The fourth tarsal subsegment always bears spines, and the spines are
MEM. AMER. ENT. SOC, 24
36 THE SATYRIDAE
present on the third and fourth subsegments in Gnophodes and Par-
antirrhoea, on the second, third and fourth in Melanitis and Hipio.
The midtibia is greater than twice as long as the proximal midtarsal
subsegment, is smooth dorsad and bears moderately well developed
tibial spurs but no spine at the dorsal, distal end. Relative lengths of
legs are shown in Fig. 20.
The forewing cell varies from just over half to nearly three-fifths
as long as the forewing costa and is somewhat excavate at ni2-m3.
The forewing radial veins arise in three branches. Veins Rs and Mi
are usually well separated at their origins, vein M- arises much closer
to Mi than to M3 and Cui usually arises midway between M:s and Cui;,
although Cui arises noticeably nearer Cu^ in Melanitis. The forewing
veins are little inflated; any inflation is restricted to the base of Sc.
The hindwing cell is more or less rounded distad and is two-fifths
to half the length of the wing measured to the end of M3. The hind-
wing vein Sc + Ri is longer than 3A. Hindwing vein M2 arises mid-
way between Mi and M3, and veins M3 and Cui are well separated at
their origins.
The pattern is similar to that of the Antirrhini with a shaded
brown and orange upper surface and cryptic coloration on the under
surface with a brown ground color and fine, dark striations.
Genera Included in the Melanitini
Cyllogenes Butler, 1868b: 6. Type-species: Melanitis suradeva Moore, by
monotypy.
Gnophodes Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-
1852: 363. Type-species: Gnophodes parmeno Westwood, designated
by Butler, 1868a: 194.
Hipio Hubner, [1819] ( 1816-1826): 56. Type-species: Papilio constantia Cra-
mer, designated by Butler, 1867a: 279.
Melanitis Fabricius, 1807: 282. Type-species: Papilio leda Linne, designated
by Butler, 1868a: 194.
= Cyllo Boisduval, 1832: 140. Type-species: Papilio leda Linne,
designated by Scudder, 1875a: 151.
Parantirrhoea Wood-Mason, 1880: 248. Type-species: Parantirrhoea mar-
shall ii Wood- Mason, by monotypy.
Genus MANATARIA Kirby, 1908
(Figures 41-47)
Manataria Kirby, 1908: 57. Type-species: Tisiphone hercyna Hubner, by
monotypy.
LEE D. MILLER
37
0.4 x ML, FL
ML
HL
IxML
W//////A
^MMMM:- ssHxSg
V//////////A
■:■:■:■:-:■:■:■:■:':■:■:::-::-:-:■:•:-:-:■;■;]
I
41
Manataria
Figs. 41-47. Manataria hercyna (Hiibner). 41. Relative lengths of the
femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light
stippling) and hindlegs (HL, heavy stippling). The ML value is unity. The
top bar represents the measurements obtained from the male, the bottom bar
those from the female. 42. $ venation. 43. palpus. 44. £ foreleg. 45. 9
foretarsus. 46. midleg. 47. bifid tarsal claw.
This South American genus has some morphological and pattern
features which connect it to the Neorina-series of the Lethini. If this
were its position, Manataria would be the only Neotropical lethine
and not particularly closely related to the few Nearctic members of
the tribe. The bifid tarsal claws, on the other hand, connect the pres-
ent genus with the sympatric Antirrhini (although this is through the
Paleotropical Melanitini), but in most other respects Manataria has
little in common with the antirrhines. The pattern is quite typical of
the Neorina-series, even to the basad displacement of the ocellus in
hindwing space Rs-Mi of the under surface. Were it not for the vast
geographical isolation and the bifid tarsal claws I would place Mana-
taria in the Lethini, perhaps in a monotypic series, but it seems more
advisable to call attention to this aberrant genus here without assign-
ing it to a tribe, pending further information. Manataria is char-
acterized as follows:
MEM. AMER. ENT. SOC, 24
38 THE SATYRIDAE
The eyes are naked. The antennae are about half the length of
the forewing costa; the antennal club is poorly developed and less
than twice as thick as the shaft. The third segment of the palpus is
just under one-third the length of the second. The hairs of the second
palpal segment are not as long as the segment is wide.
The male foreleg is well developed with the tibia slightly longer
than the femur and a monomerous, unspined tarsus. The female
foreleg is also not significantly miniaturized with a pentamerous, un-
clubbed tarsus which is spined on the second, third and fourth sub-
segments. The midleg is slightly longer than the hindleg (Fig. 41).
The midtibia is just less than twice as long as the proximal midtarsal
subsegment, is slightly spinose dorsad, bears no spine at the dorsal,
distal end and has well developed tibial spurs. The posttarsal claws
are bifid. The relative lengths of the legs are shown in Fig. 41.
The forewing cell is produced at the origin of Ms and about half
as long as the forewing costa. The forewing radial veins arise in
three branches from the cell, and Rs and Mi are connate. Vein M2
arises nearer Mi than M3, and Cut arises slightly nearer M3 than CU2.
Forewing veins Sc and the cubital stem are slightly inflated; 2A is not.
The hindwing cell is produced at the origin of Ms and is slightly
more than two-fifths the length of the wing measured to the end of
M3. Veins Sc + Ri and 3A are of about the same length, and M 3
and Cui are separate, but approximate, at their origins. Vein M 2
arises much nearer Mi than Ms.
The upper surface is dark brown with a transverse yellow spot-
band on the forewing outside the cell. Below the pattern of the upper
surface is repeated; in addition, there are pale scrawlings at the apex
of the forewing and all over the hindwing and ocelli are developed in
space M1-M2 of the forewing and in all spaces from Rs-Mi to Cu 2 -2A
of the hindwing. The ocellus in hindwing space Rs-Mi is displaced
basad as in most of the Lethini.
Subfamily ELYMNI1NAE Herrich-Schaffer, 1864
Eurytelidae Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-
1852: 403 (in part).
Elymniina Herrich-Schaffer, 1864: 124.
= Enodiinae Clark, 1947: 149.
= Lethinae Clark, 1948: 77.
LEE D. MILLER 39
The Elymniinae are the now dominant primitive satyrid subfamily
and apparently provided the source for the Satyrinae, Eritinae and
probably the Ragadiinae — although the origin of the latter subfamily
is still a problem. In some respects the elymniines may be connected
to, and may have arisen from, the Melanitini (Biinae) through such
genera as Ptychandra and Samanta (Lethini), as discussed under the
melanitines.
Although not all Elymniinae show this characteristic and some
that are not elymniines do, most satyrids with the hindwing veins M3
and Cui connate or closely approximate at their origins are probably
referable to this subfamily. The form of the hindwing cell is also
distinctive: it is usually greatly produced marginad at the origin of
vein M3, although this is not true of some Elymniini because of dis-
placement of the venation by the androconial patch. The presence of
androconial patches in specific locations is characteristic of the tribes.
With the exception of a few lethine genera of the Pararge-series, the
forewing cell is never more than half the length of the wing, in marked
contrast with the preceding three subfamilies where the cell is seldom
Table 5
Some of the diagnostic characters of the tribes of the Elymniinae.
Character Lethini Zetherini Elymniini Mycalesini
Tibial spurs .. Present Present Absent Present
Eyes Hairy, except ..Naked Naked Naked or hairy
in Neorina-
series
Forewing
vein 2A .... Little, if at Not inflated .... Not inflated .... Subquadrate
all, inflated basal infla-
tion, except
two genera
Forewing me-
dian veins .. M 2 arises M L > arises much M-, arises much IVL arises much
nearer M x nearer M^_ nearer M 1 nearer M t
than M 3 than M 3 than M 3 than M 3
Forewing cell Rounded, ca. .. Slightly exca- .. Strongly exca- .. Strongly exca-
1 2 length vate, <1 2 vate, <1 2 vate, ca. 1/2
of forewing length of length of length of
forewing forewing forewing
MEM. AMER. ENT. SOC, 24
40 THE SATYRIDAE
much less than three-fifths the length of the forewing costa. The male
foreleg is moderately well developed in all genera and is characteristi-
cally tapered to a rather sharp point.
The metropolis of the Elymniinae is the Paleotropical area, and
three of the four tribes are found more or less exclusively there. The
other tribe, the Lethini, reaches its greatest development in the Indo-
Malayan region but has representatives in Africa, the Palearctic and
two genera in North America.
The tribes of the Elymniinae all more or less grade into one an-
other. They are the Lethini, Zetherini, Elymniini and Mycalesini.
It is difficult to determine which of the tribes is primitive and which
is derived, if any are, but the A eropetes-series of the Lethini are defin-
itely primitive, whatever the condition of the other members of the
tribe; the other tribes have about equal status. A key to the tribes
of the Elymniinae is given below and an analysis of them in Table 5.
Key to the Tribes of the Elvmniinae
1. Tibial spurs absent (Fig. 96) Elymniini
Tibial spurs present (Fig. 53, for example) 2
2. Forewing vein 2A inflated at base to a subquadrate thickening (Figs. 99,
115, for example) most Mycalesini
Forewing vein 2A, if thickened, not subquadrate at base 3
3. Eyes hairy; forewing cell frequently rounded distad (Fig. 49. for example)
Lethini (except Neoriiia-series )
Eyes naked; forewing cell frequently excavate (Fig. 90, for example) .. 4
4. Base of forewing vein Sc about three times the thickness of any other
stalk (Fig. 106) Mycalesini (Orsotriaena, Bletogona)
Bases of forewing veins little inflated, Sc seldom over twice the thickness
of any other vein 5
5. Forewing vein M., arising much nearer M x than M 3 (mj-m. one-fourth or
less as long as m 2 -m 3 ; Figs. 81, 86, 87) Zetherini
Forewing arising nearer, but not much nearer, M x than M 3
Lethini (Neorina group)
Tribe Lethini Clark, 1948
(Figures 48-80)
Enodiinae Clark, 1947: 149.
Lethinae Clark, 1948: 77.
"Lethinae" is used in preference to the older "Enodiinae" for a
variety of reasons. Enodia is generally considered at best a subgenus
LEE D. MILLER 41
of Lethe, but this is an invalid reason for the suppression of a family-
group name, according to Art. 40 of the International Code of Zoo-
logical Nomenclature (1961). Part (a), however, of the same Ar-
ticle states, "If a family-group name, changed before 1961 because
of such synonymy, has won general acceptance, it is to be maintained
in the interests of stability." This is precisely applicable to the ques-
tion at hand. Dos Passos (1964: 99) uses the collective name
"Lethinae" in preference to the older "Enodiinae". To now replace
Lethinae (or in this instance, "Lethini") with Enodiinae (Enodiini)
would not serve the interests of stability of nomenclature. Lethe is
further to be preferred, all other things being equal, inasmuch as this
genus is recognized throughout much of the world; Enodia, where it
is even recognized today, is known only in the eastern part of North
America.
The Lethini may be the focal point of much of the evolution of
the higher categories of the Satyridae. As mentioned earlier in the
discussion of the Melanitini, there are some vaguely but intriguingly
indicated connections between that group and the lethines through
such lethine genera as Ptychandra and Samanta. The venation of
the former genus, while highly modified by androconial displacement
of the forewing veins, is suggestive of the pattern in the melanitines
(Fig. 57). Samanta, while a perfectly typical lethine from almost
all other aspects, does show the bifid tarsal claws which characterize
the Melanitini and the Neotropical aberrant genus Manataria. With-
in the Elymniinae, too, some interesting gradations may be seen. The
line separating the Zetherini from the Lethini is a thin one, based
chiefly on the relative positions of some veins and naked versus hairy
eyes, respectively. The zetherines, in turn, have many characteristics
in common with the Elymniini, as will be shown below. The genus
Mandarinia is a puzzle: the distribution of androconia suggest that it
is a mycalesine, but the lack of inflation of the forewing vein 2A and
the distally rounded forewing cell are lethine characteristics: this genus
is provisionally placed in the Lethini (Fig. 73). The African mem-
bers of the A eropetes-series show yet another relationship. If one
considers these butterflies and members of the satyrine tribe Dirini
together, one can almost obtain a graded series from perfectly good
Lethini to typical Satyrinae. This is probably convergence, as will
be shown later.
MEM. AMER. ENT. SOC, 24
42 THE SATYRIDAE
The Lethini are generally characterized by the hairy eyes which
separate them from the Elymniini and the zetherines, the presence of
tibial spurs (a characteristic of the Elymniini is the lack of tibial
spurs) and by the more or less rounded forewing cell, never exca-
vate as in the elymniines and most mycalesines. Most Lethini show
a characteristic basad displacement (diastoma) of the hindwing ocel-
lus in space M1-M2. There are a number of generic series, however,
so it is difficult to distinguish all lethines by a single set of criteria.
The A eropetes-series, consisting of two genera, is confined to
South Africa and characterized by the large size, extremely primitive
pattern (Schwanwitsch, 1924), hairy eyes, connate hindwing veins
Mi and Cui, extremely spiny midtibia and lack of androconial patches
(Figs. 67-72).
The Lethe-series, sensu stricto, has twenty-five nomenclatorially,
though not necessarily biologically, valid genera, twenty-two of which
are Indo-Australian or sub-Palearctic, one East African and two North
American. These insects are medium to large-sized, have a rather
primitive pattern on the under surface, though not necessarily on the
upper, hairy eyes, connate to separate hindwing veins M3 and Cui,
dorsally smooth to only slightly spiny midtibia and frequently with
androconial patches in the usual positions (Figs. 49-59, 66).
The Pararge-series has nine Palearctic genera which differ from
the Lethe-series chiefly in the length of the forewing cell: in all other
groups the cell is about half the length of the wing, whereas in Par-
arge and its allies it is about three-fifths as long ( Figs. 60-65 ) .
The Mandarinia-series is monotypic, and is characterized by its
Mycalesis-iike androconial area and is found exclusively in China
(Figs. 73-76).
The N eorina-series is composed of four Indo-Australian genera.
These butterflies are characterized by their large size, highly modified
pattern, naked eyes, spiny midtibia and lack of androconia (Figs.
77-80).
The Lethini are characterized as follows:
The eyes are hairy in all members except those of the Neorina-
series, which have naked eyes. The antennae are generally from two-
fifths to nearly half as long as the forewing costa, but the antennae
are slightly longer than half the costa in such genera as Hermias,
Tansima and Pararge. The antennal club is moderately well devel-
LEE D. MILLER
43
0.4xML, FL
I J
[
__
__
»»
IZI
r
V////A
,A
J
*m
ML | HL
Aeropetes series
Lethe 'series, s. s.
xML,
i
*Sj :
w
Z?
Pararge
i
_........:..:..^_. ; _._.w ..i~. — J
h
..._j
Mandarinia I series
^^
z zz
e^%
:.,.i
. ..■.■.■■.■ ■ ■ ■. ■■ ■■ ■ ■ ■■■ '.'.■.■.;.' ■'.
: : : : : :v:': : : : : : :':^
Neorina series
k : : : : :j : : : : : : : : : :
Aeropetes
Paralethe
Lethe
Aphysoneura
Satyrodes
Enodia
Zophoessa
Ptychandra
Pararge
Orinoma
Kirinia
Mandarinia
Neorina
Anadebis
Fig. 48. Elymniinae: Lethini. Relative lengths of the femur + tibia +
tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and
hindlegs (HL. heavy stippling) of selected genera. In all instances the ML
value is unity. The top bar for each genus represents the measurements ob-
tained from males, the bottom bar those from females.
MEM. AMER. ENT. SOC, 24
44 THE SATYRIDAE
oped, occupying the distal quarter of the antenna and inflated two to
three times the thickness of the shaft. The relative lengths of the
second and third segments of the palpus are highly variable within
the tribe: the third segment is less than one-sixth the length of the
second in Crebeta and between one-third and one-half in Hermias; in
most genera the third palpal segment is one-fourth to one-fifth the
length of the second. The length of the hairs of the second segment
of the palpus is also variable: the hairs are shorter than the segment
is wide in Hermias, but four times as long as the segment is wide in
such genera as Magula and Lopinga; in most genera the hairs are
about three times as long as the second segment is wide.
The development of the male foreleg is also variable, although
the leg itself is relatively unminiaturized. There is a great variation
in the relative length of the forefemur and tibia, and the tarsus has
one to three subsegments and is usually unspined, although there are
many tarsal spines in such genera as Ethope and Paralethe. The
female foreleg is likewise well developed and slightly clubbed at the
end of the tarsus. The tarsus is always pentamerous with spines on
at least the second, third and fourth subsegments (also on the first
subsegment in such genera as Lethe and Ptychandra). The mid-
and hindlegs are of about the same length. The midtibia is always
less than twice the length of the first midtarsal subsegment. In the
South African Aeropetes and Paralethe the midtibia is extremely
spinose dorsad (Fig. 71 ), but the midtibia is smooth in other genera,
such as Lethe and Satyrodes (Fig. 53). The tibial spurs are always
present and well developed, but there is no spine at the dorsal, distal
end of the midtibia. The relative lengths of the legs are shown in
Fig. 48.
The following cell is square-cut or rounded distad, never excavate
as in other elymniine tribes, and about half as long as the forewing
costa (Fig. 49, for example) — but this characteristic is by no means
universal. In Ptychandra the cell is only a third the length of the
wing (Fig. 57), and in most members of the Pararge-series the cell
is about three-fifths the length of the costa ( Figs. 60, 64, 65 ) . The
forewing radial veins always arise in three branches from the cell, and
veins Rs and Mi are usually well separated at their origins, although
they may arise approximate to one another. Forewing vein M2 arises
nearer, though not much nearer, Mi than M 3 ; and Cui arises midway
LEE D. MILLER
45
Figs. 49-56. Elymniinae: Lethini. 49. Lethe europa (Fabricius), S vena-
tion. 50. L. europa, palpus. 51. L. europa, 6 foreleg. 52. L. europa, 9 fore-
tarsus. 53. L. europa, midleg. 54. Raugbia scanda (Moore), 6 venation.
55. Rhaphicera satricus (Westwood and Hewitson), £ venation. 56. Tansima
satyrina (Butler), £ venation.
between Mb and C112 in most genera, but Cui arises nearer Cu- in
several genera (e.g., Lethe, Fig. 49, Ptychandra, Fig. 57, and Pararge,
Fig. 60). Forewing vein Sc is often somewhat inflated, the cubital
stem seldom inflated, although it is much inflated in Ptychandra, and
2 A is never much swollen.
The hindwing cell is produced at the origin of vein Ms and is
MEM. AMER. ENT. SOC, 24
46
THE SATYRIDAE
two-fifths to three-fifths the length of the wing measured to the end of
Ma. Hindwing vein Sc + Ri is of equal length or shorter than 3 A.
Veins M3 and Cui are usually connate, but they are well separated
in such aberrant genera as Mandarinia, Ptychandra and Rhaphicera.
Vein Mi usually arises nearer Mi than Ms, though not much nearer.
In Aeropetes M 2 arises midway between Mi and M3, slightly nearer
Ma in the Pararge-sehes and much nearer Ms in Mandarinia.
The pattern is highly variable, the main unifying feature being the
diastoma of the hindwing ocelli in space M1-M2 which was mentioned
Figs. 57-59. Elymniinae: Lethini. 57. Ptychandra lorquinii Felder and
Felder, £ venation. 58. Enodia portlandia (Fabricins), 6 venation. 59. Saty-
rodes eurydice (Linne), $ venation.
earlier. This characteristic is by no means universal, though, and is
lacking in many genera, either because of the loss of many ocelli or
extensive realignment of all the hindwing spots. There are several
sites of androconial generation in the Lethini. Forewing patches are
uncommon and restricted to the area between the origin of the cubitus
and the inner margin. These forewing patches are rarely hair tufts
(Ptychandra, Fig. 57) and are more frequently small patches of mealy
scales under the origin of Cui. The hindwing androconial patches
are much more common and usually lie just outside the cell between
Ma and Cu 2 , although Mandarinia has a Mycalesis-\ike hair tuft along
vein Rs (Fig. 73). Hindwing hair tufts are commoner than mealy
patches, but both are present throughout the tribe.
LEE D. MILLER 47
Genera Included in the Lethini
A eropetes-ser'ies
Aeropetes Billberg, 1820: 79. Type-species: Papilio tulbaghia Linne, desig-
nated by Hemming, 1943: 23.
= Meneris Westwood, 1850, in Doubleday, Westwood and Hewitson,
1846-1852: 296. Type-species: Papilio tulbaghia Linne, by mono-
typy.
Paralethe van Son, 1955: 51. Type-species: Satyrus dendrophilus Trimen, by
original designation.
Lethe-series, sensu stricto
Aphysoneura Karsch, 1894: 190. Type-species: Aphysoneura pigmentaria
Karsch, by original designation.
= Rhaphiceropsis Sharpe, 1894: 336. Type-species: Rhaphiceropsis
pringlei Sharpe (= Aphysoneura pigmentaria Karsch), by original
designation.
Archondesa Moore, 1892 (1890-1893): 270. Type-species: Lethe lanaris But-
ler, by original designation.
Charma Doherty, 1886: 117. Type-species: Zophoessa baladeva Moore, by
original designation. Proposed as a subgenus of Lethe Hiibner.
= Putlia Moore, 1892 (1890-1893): 287. Type-species: Zophoessa
baladeva Moore, by original designation. Proposed to replace
Charma Doherty, wrongly believed to be preoccupied.
Choranesa Moore, 1892 (1890-1893): 270. Type-species: Lethe trimacula
Leech, by original designation.
Debis Doubleday and Hewitson, 1849, in Doubleday, Westwood and Hewitson,
1846-1852: pi. 61. Type-species: Debis samio Doubleday and Hewit-
son, by monotypy.
Dionana Moore, 1892 (1890-1893): 271. Type-species: Lethe margaretae
Elwes, by original designation.
Enodia Hiibner, [1819] (1816-1826): 61. Type-species: Enodia andromacha
Hiibner (= Papilio portlandia Fabricius), designated by Scudder, 1872:
26.
Hanipha Moore, 1880 (1880-1881): 18. Type-species: Lethe sihala Moore
(= Lethe dynsate Hewitson), by original designation. This name is
commonly misspelled "Hanifa".
Harima Moore, 1892 (1890-1893): 299. Type-species: Neope callipteris But-
ler, by original designation.
Hermias Fruhstorfer, 1912 (1912-1915): 324. Type-species: Satyrus verma
Kollar, by monotypy. Proposed as a "subgenus or species-group" of
Lethe Hiibner.
Kerrata Moore, 1892 (1890-1893): 285. Type-species: Lethe tristigmata El-
wes, by original designation.
Kirrodesa Moore, 1892 (1890-1893): 237. Type-species: Debis sicelis Hew-
MEM. AMER. ENT. SOC, 24
48
THE SATYRIDAE
Figs. 60-72. Elymniinae: Lethini. 60. Pararge aegeria (Linne), £ vena-
tion. 61. P. aegeria, palpus. 62. P. aegeria, $ foreleg. 63. P. aegeria, 2
foretarsus. 64. Crebeta deidamia (Eversmann), $ venation. 65. Lopinga
dumetorum (Oberthiir), venation of £ forewing. 66. Aphysoneura pigmentaria
Karsch, venation of £ forewing. 67. Aeropetes tulbaghia (Linne), cS venation.
68. A. tulbaghia, palpus. 69. A. tulbaghia, £ foreleg. 70. A. tulbaghia, 2
foretarsus. 71. A. tulbaghia, midleg. 72. Paralethe dendrophiius (Trimen),
£ venation.
itson, by original designation.
Lethe Hiibner, [1819] (1816-1826): 56. Type-species: Papilio europa Fabri-
cius, by monotypy.
= Tanaoptera Billberg, 1820: 79. Type-species: Papilio europa Fab-
LEE D. MILLER 49
ricius, designated by Hemming, 1933: 199.
Magula Fruhstorfer, 1912 (1912-1915): 313. Type-species: Zophoessa jala-
urida deNiceville, designated by Hemming, 1935: 1. Preoccupied by
Magula Scudder (Megerle MS.), 1882, but no replacement name has
been proposed.
Nemetis Moore, 1892 (1890-1893): 237. Type-species: Papilio minerva Fab-
ricius, by original designation.
Neope Butler, 1867d: 166. Type-species: Lasiommata (?) bhadra Moore,
designated by Butler, 1868b: 112.
= Enope Moore, 1857: 228. Type-species: Lasiommata (?) bhadra
Moore, designated by Moore, 1892 (1890-1893): 299. Preoc-
cupied by Enope Walker, 1854.
= Blanaida Kirby, 1877: 699. Type-species: Lasiommata (?) bha-
dra Moore, by original designation. Proposed to replace Enope
Moore.
Ninguta Moore, 1892 (1890-1893): 310. Type-species: Pronophila schrenkii
Menetries, by original designation.
= A randa Fruhstorfer, 1909: 134. Type-species: Pronophila schrenkii
Menetries, by original designation.
Patala Moore, 1892 (1890-1893) : 305. Type-species: Zophoessa yama Moore,
by original designation.
Placilla Moore, 1892 ( 1890-1893) : 253. Type-species: Lethe christophi Leech,
by original designation.
Ptychandra Felder and Felder, 1861: 304. Type-species: Ptychandra lorquinii
Felder and Felder, by monotypy.
Rangbia Moore, 1892 (1890-1893): 232. Type-species: Debis scanda Moore,
by original designation.
Samanta Moore. 1880: 166. Type-species: Myca'esis malsara Moore, by
original designation.
Satyrodes Scudder, 1875b: 242. Type-species: Papilio eurydice Linne. by
original designation.
= Argus Scopoli, 1777: 432. Type-species: Papilio eurydice Linne.
designated by Scudder, 1872: 27. Preoccupied by Argus Bohadsch.
1761.
Sinchula Moore. 1892 (1890-1893): 275. Type-species: Debis sidonis Hew-
itson, by original designation.
Tansima Moore, 1881: 305. Type-species: Lethe satyrina Butler, by mono-
typy.
Zophoessa Westwood. 1851. in Doubleday, Westwood and Hewitson, 1846-
1852: 362. Type-species: Zophoessa sura Westwood. by monotypy.
Pararge-series,
Chonala Moore, 1893 (1890-1893): 14. Type-species: Debis (Tansima)
masoni Elwes, by original designation.
MEM. AMER. ENT. SOC, 24
50 THE SATYRIDAE
Crebeta Moore, 1893 (1890-1893): 11. Type-species: Pararge deidamia Evers-
mann, by original designation.
Kirinia Moore, 1893 (1890-1893): 14. Type-species: Lasiommata epimen-
ides Menetries, by original designation.
Lasiommata Westwood, 1841: 65. Type-species: Papilio megera Linne, desig-
nated by Scudder. 1875a: 202.
= Amecera Butler, 1867d: 162. Type-species: Papilio megera Linne,
designated by Butler, 1868b: 123.
Lopinga Moore, 1893 (1890-1893): 11. Type-species: Pararge dumetorum
Oberthiir, by original designation.
Orinoma Gray, 1846: 14. Type-species: Satyrus (?) damaris Doubleday {nee.
Gray), by monotypy.
Pararge Hiibner, [1819] (1816-1826) : 59. Type-species: Papilio aegeria Linne,
designated by Butler, 1868a: 195. This name has been frequently mis-
spelled as "Pararga" or "Parage" in the literature.
Rhaphicera Butler. 1867d: 164. Type-species: Lasiommata satricus Westwood
and Hewitson, designated by Butler. 1868b: 158.
Tatinga Moore, 1893 (1890-1893): 5. Type-species: Satyrus thibetanus Ober-
thiir, by original designation.
Mandarinia-series
Mandarinia Leech, 1892: 9. Type-species: Mycalesis regalis Leech, by original
designation.
Neorina-series
Ethope Moore, 1865: 770. Type-species: Mycalesis (?) himachala Moore, by
monotypy.
= Theope Moore, 1857: 234. Type-species: Mycalesis (?) hima-
chala Moore, by original designation. Preoccupied by Theope
Doubleday and Hewitson, 1847.
= Anadebis Butler, 1867c: 50. Type-species: Mycalesis (?) hima-
chala Moore, by monotypy.
Euploeamima Holland, 1887: 113. Type-species: Zethera diademoides Moore,
by original designation.
Hermianax Fruhstorfer, 1912 (1912-1915): 326. Type-species: Neorina lati-
picta Fruhstorfer (= Neorina lowi Doubleday and Hewitson), by origi-
nal designation. Proposed as a subgenus of Neorina Westwood.
Neorina Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852:
369. Type-species: Neorina hilda Westwood. by original designation.
Figs. 73-87. Elymniinae: Lethini (Figs. 73-80) and Zetherini (Figs. 81-
87). 73. Mandarinia regalis (Leech), 6 venation. 74. M. regalis, palpus.
75. M. regalis, 9 foretarsus. 76. M. regalis, midleg. 77. Neorina hilda West-
wood, 6 venation. 78. N. hilda, $ foreleg. 79. N. hilda, 2 foretarsus. 80.
LEE D. MILLER
51
N. Hilda, midleg. 81. Zethera pimplea (Erichson), 6 venation. 82. Z. pim-
ple a, palpus. 83. Z. pimplea, 6 foreleg. 84. Z. pimplea, 2 foretarsus. 85.
Z. pimplea, midleg. 86. Amechania incerta Hewitson. i venation. 87. Cal-
large sagitta (Leech). 6 venation.
MEM. AMER. ENT. SOC. 24
52
THE SATYRIDAE
Tribe Zetherini, new tribe
(Figures 81-88)
The zetherines have been considered part of the Elymniini by
such authors as Gaede (1931), and while these two tribes share such
features as close forewing veins M- and Cui, they are abundantly
distinct. Perhaps the closest relatives of the zetherines are certain
of the more primitive lethines such as Aeropetes, but the differences
discussed below also serve to separate these two tribes. The pattern
of the zetherines is highly aberrant: several species are white with
brown interveinal markings and either no ocelli or poorly developed
ones, whereas Zethera pimplea (Erichson) looks like the Nearctic
0.4xML H
1 WZ/
Zethera
Amechania
Callarge
Fig. 88. Elymniinae: Zetherini. Relative lengths of the femur + tibia +
tarsus of the forelegs (FL, diagonal lines), midlegs (ML. light stippling) and
hindlegs (HL, heavy stippling) of the genera. In all instances the ML value
is unity. The top bar for each genus represents the measurements obtained
from males, the bottom bar those from females.
Limenitis weidemeyeri (Edwards) (Nymphalidae: Limenitinae), a
black species with a wide white central band crossing both wings.
The absence of inflation of the forewing veins serves to distinguish
this tribe from the Mycalesini, while the spiny midtibia and the well
developed tibial spurs separate the Zetherini from the Elymniini.
Most Lethini, with the exception of the Neorina-series, have hairy
eyes, whereas all zetherines have naked eyes. The relative position
of the forewing veins M.i, Cui and Cu 2 serve to distinguish members
of this tribe from the Neorina-series of the Lethini. All the members
of the Zetherini are Indo- Australian. The zetherines are character-
ized as follows:
The eyes are naked. The antennae are just over two-fifths the
LEE D. MILLER 53
length of the forewing costa. The antennal club is weakly developed
and gently tapered, never more than twice the thickness of the shaft.
The relative lengths of the second and third segments of the palpus
are variable: in Collar ge the third segment is about one-seventh as
long as the second, whereas in Zethera it is one-third as long. The
hairs of the second segment of the palpus are always shorter than the
segment is wide.
The male foreleg is well developed with the femur longer than the
tibia and a mono- to trimerous tarsus which may or may not bear
spines. The female foreleg is also well developed with a pentamerous,
slightly clubbed tarsus bearing spines on the first four subsegments.
The hindlegs are equal to or slightly longer than the midlegs (Fig.
88). The mid-tibia is long twice to two and a half times as
long as the first midtarsal subsegment — and the midtibia is spiny
dorsad, bearing moderately well developed tibial spurs, but no spine
at the dorsal, distal end.
The forewing cell is slightly excavate and two-fifths to half as long
as the forewing costa. The forewing radial veins arise from the cell
in three branches, and veins Rs and Mi are approximate, but not
connate, at their origins. Vein M2 arises much nearer Mi than M. f ,
and Cui arises much nearer M3 than Cu 2 . None of the forewing
veins are much inflated basally.
The hindwing cell is strongly produced at the origin of M3 and
is one-third to half the length of the wing measured to the end of
M3. Hindwing vein Sc + Ri is longer than 3 A. Veins M3 and Cui
are connate, or nearly so, and M- arises nearer (or in some cases,
much nearer) Mi than Ma.
The pattern of all the species is aberrant. Callarge is cream-
colored with dark veins. Amechania is whitish with dark veins, some
dark interveinal markings and a few ocelli. Zethera is blackish-
brown with broad discal white bands on all wings. There are no
defined androconial patches on any zetherine.
Genera Included in the Zetherini
Amechania Hewitson, 1861 (1856-1876): 87. Type-species: Amechania in-
certa Hewitson, by monotypy.
Callarge Leech. 1892: 57. Type-species: Zethera sagitta Leech, by original
designation.
MEM. AMER. ENT. SOC, 24
54
THE SATYRIDAE
Zethera C. Felder, 1861: 26. Type-species: Cynthia pimplea Erichson, by
monotypy.
Tribe Elymniini Herrich-Schaffer, 1864
(Figures 89-97)
Eurytelidae Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-
1852: 403 (in part).
Elymniina Herrich-Schaffer, 1864: 124.
The Elymniini are a homogeneous group and show some affinities
within the subfamily. There is little to connect the elymniines with
the Lethini directly, but the positions of certain of the veins show a
definite relationship between these butterflies and the zetherines,
which in turn do relate to the Lethini. The deeply excavate forewing
cell of these insects is approached in the Mycalesini, and many of the
latter groups have androconial patches placed very like those which
characterize the Elymniini.
0.4xML, FL
Elymnias
Elymniopsis
Melynias
Mimadelias
Biudsa
Fig. 89. Elymniinae: Elymniini. Relative lengths of the femur + tibia
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML
value is unity. The top bar for each genus represents the measurements ob-
tained from males, the bottom bar those from females.
This tribe may be distinguished from all others within the sub-
family by the lack of tibial spurs in all species. Many of the genera
also have extremely short cells on both wings, but there are a few
exceptions.
LEE D. MILLER 55
All elymniines are Paleotropical, most being found in the Indo-
Australian region, but a couple of species range as far west as West
Africa. The Elymniini are characterized in the following paragraphs.
The eyes are naked. The antennae are about two-fifths as long
are the forewing costa, although they may be as much as 0.46 times
as long as the costa. The antennal club is very gentle and seldom
more than twice the width of the shaft, except in the genus Melynias
where it is over twice the thickness of the shaft. The third segment
of the palpus is from one-fifth {Melynias) to one-third (Dyctis) the
length of the second segment. The hairs of the second palpal seg-
ment are about as long as the segment is wide.
The male foreleg is well developed with the femur longer than
the tibia and with a monomerous, unspined tarsus. The female fore-
leg is also well developed with a pentamerous, unclubbed tarsus
bearing a double set of spines on the third and fourth subsegments
(also a double set on the second subsegment in Elymniopsis) . The
mid- and hindlegs are of about the same length (Fig. 89). The
midtibia is generally less than twice the length of the first midtarsal
subsegment and is smooth dorsad, without tibial spurs or a spine at
the dorsal, distal end.
The forewing cell is usually deeply excavate (Figs. 90-92) and
is always less than half the length of the forewing costa. Frequently
the cell is less than a third as long as the costa. The forewing radial
veins arise in three branches from the cell, and Rs and Mi are well
separated at their origins. Vein M 2 arises much nearer Mi than M.3,
and Cui is almost connate with M3, never near Cu-. Forewing vein
Sc is usually inflated basad; the other stems are not.
The hindwing cell is more or less square-cut because of the dis-
placement marginad of Mi and is from one-third to almost half as
long as the wing measured to the end of M3. Hindwing vein Sc + Ri
is always much shorter than 3A. Veins M3 and Cui are always
connate, or nearly so, and M2 arises much nearer Mi than M3.
The pattern is quite variable and has been considered a mimetic
one. Many of the species resemble various Danaidae, particularly of
the genus Euploea. For a description of most of the species see
Moore, 1893 (1890-1893).
MEM. AMER. ENT. SOC, 24
56
THE SATYRIDAE
Figs. 90-97. Elymniinae: Elymniini. 90. Elymnias hypermnestra (Linne),
S venation. 91. Bruasa penanga (Westwood and Hewitson), S venation.
92. Dyctis agondas Boisduval, 6 venation. 93. E. hypermnestra, palpus. 94.
E. hypermnestra, S foreleg. 95. E. hypermnestra, 9 foretarsus. 96. E. hy-
permnestra, midleg. 97. Elymniopsis Use Hemming, palpus.
Genera Included in the Elymniini
A grusia Moore, 1893 (1890-1893): 144. Type-species: Melanitis esaca West-
wood and Hewitson, by original designation.
Bruasa Moore, 1893 (1890-1893): 144. Type-species: Melanitis penanga
Westwood and Hewitson, by original designation.
Didonis Hiibner, [1819] (1816-1826): 17. Type-species: Papilio vitellia
Cramer, designated by Scudder, 1875a: 156.
LEE D. MILLER 57
Dyctis Boisduval. 1832: 138. Type-species: Dyctis agondas Boisduval, by
monotypy.
Elymnias Hiibner, 1818 (1818-1825): 12. Type-species: Elymnias jynx Hiib-
ner (= Papilio hypermnestra Linne), designated by Hemming, 1943:
24. Scudder (1875a: 162) designated Papilio lais Fabricius as type,
based on Elymnias Hiibner, [1819] (1816-1826): 37. This designation
was invalidated when the appropriate sections of the "'Zutrage" were
found to be older than those of the "Verzeichniss".
Elymniopsis Fruhstorfer. 1907: 171, 173-174. Type-species: Papilio phegea
Fabricius (= Elymniopsis Use Hemming), designated by Hemming,
1943: 24.
Melynias Moore, 1893 (1890-1893): 144. Type-species: Papilio lais Cramer
(= Papilio nesaea Linne), by original designation.
Mimadelias Moore, 1893 (1890-1893): 144. Type-species: Elymnias vasu-
deva Moore, by original designation.
Tribe Mycalesini, new tribe
(Figures 98-119)
The mycalesines show rather close relationships to both the
Lethini and the Elymniini. Such mycalesine genera as Orsotriaena
and Bletogona lack the characteristic inflation of the veins, but do
have the excavate forewing cell and the distinctive androconial
patches, whereas the lethine Mandarinia regalis (Leech) has the
androconial patch but lacks the inflation of forewing vein 2A or the
excavate forewing cell. The excavate forewing cell and the presence
in some genera of a hair tuft on the upper surface of the hindwing
at the anterior margin of the cell strongly suggests a relationship
between the Mycalesini and the Elymniini.
The characteristic subquadrate shape of the basal inflation of
forewing vein 2A serves to distinguish the vast majority of all
mycalesines. In the Hypocystini (Satyrinae) this inflation of 2A
is suggested, but in that tribe the inflation is in the form of a rounded
knot, rather than a subquadrate thickening, as in the present tribe.
The Mycalesini are general throughout the Old World tropics and
form a homogeneous group. There is some regional correlation of
genera in the tribe, and for that reason they are arranged geo-
graphically here. It is interesting to note that although hairy or
naked eyes do not characterize the tribe, all mainland African Myca-
lesini have naked eyes, all those from Madagascar have hairy eyes
and both states are found in the Indo-Australian genera. This
MEM. AMER. ENT. SOC, 24
58
THE SATYRIDAE
0.4 XML, FL
Mycalesis
Satoa
Orsotriaena
Bletogona
Bicyclus
Dichothyris
Admiratio
Heteropsis
Masoura
Fig. 98. Elymniinae: Mycalesini. Relative lengths of the femur + tibia
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and
hind legs (HL, heavy stippling) of selected genera. In all instances the ML
value is unity. The top bar for each genus represents the measurements ob-
tained from males, the bottom bar those from females.
situation will be discussed further in the section on "Evolution and
Zoogeography." The Mycalesini are characterized as follows:
The eyes are naked or hairy: the Indo- Australian species show
both conditions, those from mainland Africa all have naked eyes
and those from Madagascar have hairy eyes. The antennae are from
two-fifths to half the length of the forewing costa. The antennal club
is moderately well developed, occupying the distal one-third to one-
fifth of the antenna and thickened from less than twice ( Orsotriaena )
to more than three times (such genera as Dalapa and Martanda) the
thickness of the shaft. The third segment of the palpus is usually
one-third to one-fifth the length of the second segment, but the third
LEE D. MILLER 59
segment is more than half as long as the second in Dalapa. The hairs
of the second palpal segment are generally from as long as the seg-
ment is wide to twice the width of the segment.
The male foreleg is well developed with the femur longer than
the tibia and one or (rarely) two unspined or weakly spined tarsal
subsegments. The female foreleg is moderately well developed with
a pentamerous, lightly clubbed (less so in genera from Madagascar,
Fig. 118) tarsus bearing spines on at least the fourth subsegment,
often also on the second and third and occasionally on the first (such
genera as Martanda) . The mid- and hindlegs are not significantly
unequal. The midtibia is less than twice as long as the first midtarsal
subsegment. The midtibia is usually unspined dorsad, although it is
weakly spined in Orsotriaena (Fig. 108), etc.; tibial spurs are always
present and well developed and there is no spine at the dorsal distal
end of the midtibia.
The forewing cell is excavate and varies little from half the length
of the forewing costa. The forewing radial veins arise in three
branches from the cell, and veins Rs and Mi arise separate, but
approximate, from the cell. Vein M2 arises much nearer Mi than
Ms, and Cui arises nearer Ma than Cu-. The pattern of basal infla-
tion of the forewing veins is characteristic, except in Orsotriaena
(Fig. 106) and Bletogona: veins Sc and the cubital stem are inflated
normally, but markedly, and 2A is inflated near its base as a sub-
quadrate knot.
The hindwing cell is produced at M:s and is one-third to half as
long as the wing measured to the end of Ms. Hindwing vein Sc + Ri
is as long as, or shorter than, 3A. Veins Ms and Cui are connate or
approximate at their origins, except in Culapa (Fig. 104) where they
are widely separated, and M2 arises much nearer Mi than M3.
The pattern is highly variable. Many genera show the greatest
development of the ocelli in forewing space Cui-Cu:>. Most of the
species are dark brown on both surfaces, hence the common name
"bushbrowns", but some of the African species are iridescent blue
or purple above and the Malgache species Masoura masoura (Hewit-
son) is white with a yellow patch on the forewing, superficially more
resembling a pierid than a satyrid. The distribution of androconia
is characteristic: there is a polished area along the inner margin of
the forewing on the under surface, and other areas of androconial
concentration are in the area between Cu-2 and 2A of the forewing
MEM. AMER. ENT. SOC, 24
60
THE SATYRIDAE
Figs. 99-108. Elymniinae: Mycalesini. 99. Mycalesis jrancisca (Stoll),
$ venation. 100. M. jrancisca, palpus. 101. M. jrancisca, S foreleg. 102.
M. jrancisca, 2 foretarsus. 103. M. jrancisca, midleg. 104. Culapa mnasicles
(Hewitson), 8 venation. 105. Mydosama juscum (Felder and Felder), $
venation. 106. Orsotriaena medus (Fabricius), 6 venation. 107. O. medus,
2 foretarsus. 108. O. medus, midleg.
and along the hindwing cell on both margins,
frequent than mealy patches in this tribe.
Hair tufts are more
Genera Included in the Mycalesini
Indo-Australian genera
Bletogona Felder and Felder, 1867 (1864-1867): 465. Type-species: Bleto-
LEE D. MILLER 61
gona mycalesis Felder and Felder, by monotypy.
Calysisme Moore, 1880 (1880-1881): 20. Type-species: Papilio drusia Cra-
mer, designated by Moore, 1891 (1890-1893): 172.
Celebina Fruhstorfer, 1898: 263. Type-species: Celebina inga Fruhstorfer,
designated by Hemming, 1935: 1.
Culapa Moore, 1878: 825. Type-species: Mycalesis mnasicles Hewitson, by
monotypy.
Dolapa Moore, 1880: 158. Type-species: Mycalesis sudra Felder, by original
designation.
Hamadryopsis Oberthiir, 1899: 17. Type-species: Hamadryopsis drusillodes
Oberthiir, by monotypy.
= Drusillopsis Fruhstorfer, 1908: 217. Type-species: Hamadryopsis
drusillodes Oberthiir, by monotypy.
Indalasa Moore, 1880: 166. Type-species: Mycalesis moorei Felder, by mono-
typy.
Jatana Moore, 1880: 164. Type-species: Mycalesis mynois Hewitson, by
monotypy.
Kabanda Moore, 1880: 168. Type-species: Mycalesis malsarida Butler, by
original designation.
Loesa Moore, 1880: 177. Type-species: Mycalesis oroatis Hewitson, by mono-
typy.
Lohora Moore. 1880: 175. Type-species: Mycalesis dexamenus Hewitson, by
original designation.
Martanda Moore, 1880: 169. Type-species: Mycalesis janardana Moore, by
original designation.
Monotrichtus Hampson, 1891: 179. Type-species: Mycalesis safitza Hewitson,
by original designation. Originally proposed as a subgenus of Myca-
lesis Hiibner.
Mycalesis Hiibner, 1818 (1818-1825): 17. Type-species: Papilio francisca
Stoll, designated by Hemming, 1937: 149. Butler, 1868a: 196, desig-
nated Papilio evadne Cramer as type, based on Mycalesis Hiibner,
[1819] (1816-1826): 55. This designation was invalidated when the
appropriate sections of the "Zutrage" were found to be older than those
of the "Verzeichniss".
= Gareris Moore, 1880: 156. Type-species: Mycalesis sanatana
Moore (= Papilio francisca Stoll), by original designation.
Mydosama Moore, 1880: 170. Type-species: Dasyomma fuscum Felder and
Felder, by original designation.
= Dasyomma Felder and Felder, 1860: 401. Type-species: Dasy-
omma fuscum Felder and Felder, by monotypy. Preoccupied by
Dasyomma Macquart, 1841.
Myrtilus deNiceville, 1891: 341. Type-species: Mycalesis (Myrtilus) mystes
deNiceville, by original designation. Proposed as a subgenus of Myca-
lesis Hiibner.
MEM. AMER. ENT. SOC, 24
62
THE SATYRIDAE
Figs. 109-119. Elymniinae: Mycalesini. 109. Bicyclus hewitsonii (Dou-
met), $ venation. 110. B. hewitsonii, palpus. 111. B. hewitsonii, $ foreleg.
112. B. hewitsonii, ? foretarsus. 113. Dichothyris sambulos (Hewitson), $
venation. 114. Heteropsis drepana Westwood, antennal club. 115. H. dre-
pana, $ venation. 116. H. drepana, palpus. 117. H. drepana, S foreleg.
118. H. drepana, 2 foretarsus. 119. Admiratio paradoxa (Mabille), $ vena-
tion.
Nasapa Moore, 1880: 176. Type-species: Mycalesis aramis Hewitson, by
monotypy.
Nebdara Moore, 1880: 173. Type-species: Mycalesis tagala Felder, by origi-
nal designation.
Nissanga Moore, 1880: 169. Type-species: Mycalesis patnia Moore, by origi-
nal designation.
Orsotriaena Wallengren, 1858: 79. Type-species: Papilio medus Fabricius ( =
Papilio hesione Cramer), by monotypy.
Pachama Moore, 1880: 165. Type-species: Mycalesis mestra Hewitson, by
monotypy.
LEE D. MILLER 63
Physcon deNiceville, 1898: 133. Type-species: Mycalesis pandoea Hopffer, by
original designation. Proposed as a subgenus of Mycalesis Hubner.
Sadarga Moore, 1880: 157. Type-species: Mycalesis gotama Moore, by origi-
nal designation.
Samundra Moore, [1890-1892] (1890-1893): 162. Type-species: Mycalesis
anaxioides Marshall and deNiceville, by monotypy.
Satoa Moore, 1880: 157. Type-species: Mycalesis maianeas Hewitson, by
monotypy.
Sevanda Moore, 1880: 174. Type-species: Satyrus duponcheli Guerin, by
original designation.
Suralaya Moore, 1880: 159. Type-species: Mycalesis oreseis Hewitson, by
monotypy.
Telinga Moore, 1880: 167. Type-species: Satyrus adolphei Guerin, by mono-
typy.
Virapa Moore, 1880: 156. Type-species: Mycalesis anoxias Hewitson, by origi-
nal designation.
African genera
Bicyclus Kirby, 1871a: 47. Type-species: Idiomorphus hewitsonii Doumet,
designated by Kirby, 1871b: 363.
= Idiomorphus Doumet, 1861: 174. Type-species: Idiomorphus
hewitsonii Doumet, by monotypy. Preoccupied by Idiomorphus
Chaudoir, 1846.
Dichothyris Karsch, 1893: 203. Type-species: Mycalesis sambulos Hewitson,
designated by Hemming, 1935: 1.
Hallelesis Condamin, 1960: 1257. Type-species: Mycalesis asochis Hewitson.
designated by Condamin, 1961: 783.
Malgache genera
Admiratio Hemming, 1964b: 137. Type-species: Smithia paradoxa Mabille,
by original designation.
= Smithia Mabille, 1880: 173. Type-species: Smithia paradoxa
Mabille, by monotypy. Preoccupied by Smithia Milne Edwards
and Haime, 1851.
Henotesia Butler, 1879: 228. Type-species: Henotesia wardii Butler, by origi-
nal designation.
Heteropsis Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-
1852: 323. Type-species: Heteropsis drepana Westwood, by monotypy.
Houlbertia Oberthur, 1916: 199. Type-species: Erebia passandava Ward, des-
ignated by Hemming, 1964a: 120.
Masoura Hemming, 1964b: 138. Type-species: Melanitis masoura Hewitson,
by original designation.
= Gallienia Oberthur, 1916: 205. Type-species: Melanitis masoura
Hewitson, designated by Hemming, 1964a: 120.
MEM. AMER. ENT. SOC, 24
64
THE SATYRIDAE
Subfamily Eritinae, new subfamily
It is obvious that the two genera in this subfamily are intermediate
between the Elymniinae (and closest to the Lethini of that subfamily)
and the Satyrinae, but perhaps the present genera should not be com-
bined into a single sub-family or tribe. Nevertheless, these genera
seem closest to one another, and I should be reluctant to erect two
monotypic taxa for them.
The female foretarsus is pentamerous and unclubbed (Figs. 125,
131) and more primitive than that of the Satyrinae. The pattern
is rather more primitive than that of most satyrines (Schwanitsch,
1924) and more closely approximates the pattern of the lethines.
The male foreleg of the genus Coelites is much reduced over the
situation in the lethines, however, and the forewing cell of Erites
is much longer than that of any elymniine.
Both genera are found on the Malay peninsula and in Indonesia.
A single tribe, the Eritini, includes all species.
Tribe Eritini, new tribe
(Figures 120-132)
The diagnostic features of the subfamily Eritinae and of the only
tribe, the Eritini, are as follows:
The eyes are naked. The antennae are about two-fifths the length
0.4* ML
Erites
Coelites
Fig. 120. Eritinae: Eritini. Relative lengths of the femur + tibia + tar-
sus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and
hindlegs (HL, heavy stippling) of the genera. In all instances the ML value
is unity. The top bar for each genus represents the measurements obtained
from males, the bottom bar those from females.
of the forewing costa. The antennal club is indistinct {Coelites,
Fig. 122) to gradual (Erites, Fig. 128) and never more than twice
as thick as the shaft. The third segment of the palpus is about
LEE D. MILLER 65
one-fifth as long as the second, and the development of the hairs of
the second segment is variable: in Coelites they are about as long as
the segment is wide, whereas in Erites the hairs are two and a half
times the width of the segment.
The male foreleg is variably developed: that of Erites is slightly
miniaturized with subequal femur and tibia and a monomerous,
unspined tarsus (Fig. 130); whereas that of Coelites is greatly
reduced, the femur longer than the tibia and the tarsus represented
only by a bump at the tip of the tibia (Fig. 124). The female foreleg
is moderately well developed with a pentamerous, unclubbed tarsus
bearing spines on the third and fourth subsegments {Erites, Fig. 131)
or on the first four subsegments (Coelites, Fig. 125). The midlegs
are approximately equal to, or slightly longer than, the hindlegs
(Fig. 120). The midtibia is just over twice as long as the first
midtarsal subsegment and is smooth dorsad with moderately well
developed tibial spurs and no spines at the dorsal, distal end.
The forewing cell is produced at the origin of Mi and one-half
{Coelites, Fig. 121) to three-fifths (Erites, Fig. 127) as long as the
forewing costa. The forewing radial veins arise in three branches
from the cell, and Rs and Mi are approximate. Vein M-- arises
slightly nearer (Erites) to much nearer (Coelites) Mi than M 3 , and
Cui arises somewhat nearer M.3 than Cuo. Only forewing vein Sc
is inflated basad.
The hindwing cell is produced at the origin of Ms and is less
than half (Coelites, Fig. 121) to three-fifths (Erites, Fig. 127) as
long as the wing measured to the end of Ms. Hindwing vein Sc + Ri
is longer than 3 A, and veins M 3 and Cui may be connate (Coelites)
to widely separated (Erites) at their origins. Vein Mi; arises slightly
nearer Mi than Ms.
Erites is an olive-brown insect above with the lines of the under
surface faintly indicated and ocelli in forewing spaces M1-M2, M2-M3
and a large one in Cui-Cu^ and hindwing spaces Rs-Mi through
CU1-CU2. On the under surface the pattern is the same but there
are dark striatums and transverse ochre lines in the medial areas of
both wings. Coelites is uniform gray-brown above with a basal
purplish sheen and a patch of oily hairs between hindwing veins 2A
and 3A. The under surface is dark brown basad, gray-brown mar-
MEM. AMER. ENT. SOC, 24
66
THE SATYRIDAE
Figs. 121-132. Eritinae: Eritini. 121. Coelites nothis Westwood and
Hewitson, $ venation. 122. C. nothis, antennal club. 123. C. nothis, palpus.
124. C. nothis, S foreleg. 125. C. nothis, 9 foretarsus. 126. C. nothis, mid-
leg. 127. Erites madura (Horsefield), S venation. 128. E. madura, antennal
club. 129. E. madura, palpus. 130. E. madura, S foreleg. 131. E. madura,
$ foretarsus. 132. E. madura, midleg.
ginad with ocelli in hindwing spaces Rs-Mi through Cui-Cul..
Genera Included in the Eritini
Coelites Westwood and Hewitson, 1850, in Doubleday, Westwood and Hewit-
son, 1846-1852: pi. 66. Type-species: Coelites nothis Westwood and
Hewitson, designated by Butler, 1868a: 195.
Erites Westwood, 1850, in Doubleday, Westwood and Hewitson, 1846-1852:
392. Type-species: Hipparchia madura Horsefield, by monotypy.
LEE D. MILLER
67
Subfamily Ragadiinae Herrich-Schaffer, 1864
Ragadiina Herrich-Schaffer, 1864: 124.
It is difficult to place the Ragadiinae in a satisfactory systematic
position, but for reasons to be enumerated below, this subfamily
appears to occupy a position intermediate between the Elymniinae
and the Satyrinae. In common with the former, hindwing veins
M:s and Cui arise from a common point on the cell, but the abortion
of the forelegs of both sexes (Figs. 136, 137) is a characteristic
shared with the Satyrinae and Coelites of the Eritinae. In other
respects, however, the ragadiines stand apart from all of the other
subfamilies.
The configuration of the hindwing cell (Figs. 134, 139) is dis-
tinctive. In males the cell is closed by narrow vestigial veins, whereas
in the females of some species the cell is completely open between the
origins of veins M2 and M 3 .
All members of the Ragadiinae are native to the Indo-Australian
region and constitute a single tribe, the Ragadiini. Rober (1892)
split this subfamily, referring Acrophtalmia to the Ypthima group
and Ragadia to the Mycalesis group, but the characteristic venation
of the hindwing cannot be ascribed merely to the convergence of two
different phyletic lines.
Tribe Ragadiini Herrich-Schaffer, 1864
(Figures 133-139)
Ragadiina Herrich-Schaffer, 1864: 124.
0.2xML
Ragadia
Acrophtalmia
Fig. 133. Ragadiinae: Ragadiini. Relative lengths of the femur + tibia
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML
value is unity. The top bar for each genus represents the measurements ob-
tained from males, the bottom bar those from females.
MEM. AMER. ENT. SOC, 24
68
THE SATYRIDAE
Figs. 134-139. Ragadiinae: Ragadiini. 134. Ragadia crisia (Geyer), $
venation. 135. R. crisia, palpus. 136. R. crisia, 8 foreleg, showing also the
trochanter and the distal portion of the coxa. 137. R. crisia, 9 foretarsus.
138. R. crisia, midleg. 139. Acrophtalmia artemis Felder and Felder, S vena-
tion.
The diagnostic features of the subfamily Ragadiinae and its only
tribe, the Ragadiini, are as follows:
The eyes are naked. The antennae are no more than half the
length of the forewing costa. The antennal club is moderately well
developed, occupies the distal one-fourth to two-fifths of the antenna
and is thickened to two and a half to three times the width of the
shaft. The third segment of the palpus is from less than a quarter
(Ragadia) to slightly less than one-third (Acrophtalmia) the length
of the second segment. The hairs of the second segment of the
palpus are shorter than the segment is wide.
The male foreleg is greatly reduced with but a bump representing
the tarsus in some species (Ragadia, Fig. 136, for example); the
tarsus is completely fused with the tibia and indistinguishable from
it in Acrophtalmia. The femur and tibia are of about the same
length. The female foreleg is also miniaturized and the tarsus clubbed
with five subsegments bearing spines on the first four. The mid- and
hindlegs are of about the same length. The midtibia is less than
twice the length of the first midtarsal subsegment, is smooth dorsad
and bears weakly developed tibial spurs, but no spine at the dorsal,
distal end. The relative lengths of the legs are shown in Fig. 133.
The forewing cell is excavate near the origin of Ml> and about
LEE D. MILLER 69
three-fifths the length of the forewing costa. The forewing radial
veins arise in two branches from the cell, and veins Rs and Mi are
nearly connate. Vein M2 arises nearer Mi than M3, and Cm arises
much nearer M.h than Cu:>. Forewing vein Sc is inflated basad, but
the other veins are not.
The hindwing cell is distinctive. It is open in some females
and at most closed by small, aberrant veins (Figs. 134, 139). The
cell is from less than half {Acrophtalmia) to more than three-fifths
(Ragadia) as long as the wing measured to the end of M3. The
relative lengths of hindwing veins Sc + Ri and 3 A vary: Sc + Ri is
longer in Acrophtalmia, and 3 A is longer in Ragadia. Veins Mm and
Cui are connate, whereas M2 arises basad of Mi and much closer to
it than to M.i.
The pattern is highly modified, either brown with a broad central
white band or with alternating brown and off-white stripes. The
ocelli are poorly developed, except in Ragadia which has a full com-
plement on the under surface. An androconial hair tuft lies on the
aberrant veins of the hindwing cell (Figs. 134, 139) and is charac-
teristic of the tribe.
Genera Included in the Ragadiini
Acrophtalmia Felder and Felder, 1861: 305. Type-species: Acrophtalmia ar-
temis Felder and Felder, by monotypy. Many authors, following Fel-
der and Felder, 1867 (1864-1867): 486, have misspelled this name
" Anophthalmia" '.
Acropolis Hemming, 1934b: 77. Type-species: Acrophthalmia (sic.) thalia
Leech, by original designation.
= Pharia Fruhstorfer, 1912 (1912-1915): 295. Type-species: Ac-
rophthalmia (sic.) thalia Leech, by monotypy. Preoccupied by
Pharia Gray, 1840.
Ragadia Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852:
376. Type-species: Euptychia crisia Geyer, by monotypy. Proposed as
a subgenus of Neonympha Hiibner.
Subfamily Satyrinae Boisduval, 1836
Satyrides Boisduval, 1836: 166; Satyridae Swainson, 1840: 86.
= Hipparchiadae Kirby, 1837: 297.
= Maniolinae Hampson, 1918: 385.
The Satyrinae are the most evolved members of the family and
MEM. AMER. ENT. SOC, 24
70
THE SATYRIDAE
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MEM. AMER. ENT. SOC, 24
72 THE SATYRIDAE
have become the dominant group throughout most of the world.
Fairly close relationships with three other subfamilies, the Elymniinae,
Eritinae and Ragadiinae, are indicated by analysis of structures, and
there are three instances of gradation between the elymniines and the
satyrines. There is probably only convergence between the Aero-
petes-series of the Lethini and the Dirini of the present subfamily.
Far more significant evolutionarily, however, are such genera as
Lamprolenis in the primitive satyrine tribe Hypocystini. These but-
terflies show certain similarities to some Mycalesini and Lethini
(Mandarinia) of the Elymniinae, particularly as regards pattern and
the placement of the androconial areas. This is probably the stock
from which the Satyrinae evolved. The Lethini are connected to
the Satyrinae through the sub-family Eritinae, as indicated in the
discussion of the eritines. This evidence seems to place the Lethini
as the closest present-day ancestral stock of the Satyrinae. The
relationship with the ragadiines is obscure and more in the nature of
an association because of the greatly reduced forelegs of both sexes.
Any satyrid with fewer than five subsegments on the female fore-
tarsus is a member of this subfamily. Even when there are five
subsegments on the female foreleg of a member of the present sub-
family, the tarsus is formed into a clublike structure totally unlike
anything found in the Haeterinae, Brassolinae, Biinae, Elymniinae or
Eritinae, but hinted at in the Ragadiinae. Comparative female fore-
tarsi of the pentamerous type are shown in Figs. 5 (Haeterinae), 15
(Brassolinae), 24, 32 (Biinae), 52, 84, 95, 102 (Elymniinae), 131
(Eritinae), 137 (Ragadiinae) and 145, 174, 200 (Satyrinae). A
greatly reduced male foreleg will serve to separate the Satyrinae from
all other satyrid subfamilies except the Ragadiinae and Coelites of the
Eritinae. None of the satyrines have the hindwing veins Ms and Cui
connate, nor the open hindwing cell of the ragadiines. The hindwing
cell, measured to the origin of vein M 3 , is greater than half as long
as the wing to the end of Ms, except in some Dirini. The antennal
club is always moderately well and frequently highly developed.
The Satyrinae are virtually cosmopolitan, with representatives
in all parts of the world where butterflies are found, except some
oceanic islands. Unlike the other subfamilies, the Satyrinae are
highly developed in temperate regions.
Ten tribes are recognized in the Satyrinae. The most primitive
LEE D. MILLER 73
tribe, the Hypocystini, contains genera which span the gap between
the other two groups. The Ypthimini-section, characterized more or
less by their smaller size, frailer appearance and an ecological prefer-
ence for grasslands, contains the tribes Ypthimini, Euptychiini, Coe-
nonymphini, Maniolini and Erebiini. The Satyrini-section, larger
and sturdier insects with woodland or scrubland preferences, is com-
posed of the tribes Dirini, Pronophilini, Satyrini and Melanargiini.
A key to the tribes of the Satyrinae is given below, and a diagnostic
chart of these tribes is provided in Table 6.
Key to the Tribes of the Satyrinae
1 . Hindwing cell half or less the length of the wing; forewing cell rounded
distad (Figs. 248, 255, 256); South Africa Dirini
Hindwing cell greater than half the length of the wing; forewing cell
square-cut or excavate 2
2. Midtibia with a long spine at the dorsal distal end (Fig. 301); Holarc-
tic most Satyrini
Midtibia without such a spine 3
3. Hindwing cell prolonged by a distad displacement of crossvein m=-m.-i
along vein M= (Fig. 261, for example) 4
Hindwing cell more or less square cut with crossvein rrb-m.! in line with
mi-rm (Fig. 309, for example) 8
4. Female foretarsus with fewer than five subsegments 5
Female foretarsus pentamerous 6
5. Third segment of palpus less than one-fourth the length of the second
(Fig. 305); Holarctic Satyrini (Oenm-series)
Third segment of palpus longer than one-fourth the length of the second
(Fig. 282); Neotropical some Pronophilini
6. Midtibia spiny; Neotropics most Pronophilini
Midtibia smooth 7
7. Neotropical species a few Pronophilini
Australian region species Hypocystini
8. White (or cream-colored) species, marbled with
black: Palearctic Melanargiini
Generally brown or orange species, if white
not marbled 9
9. Female foretarsus with fewer than five subsegments 10
Female foretarsus pentamerous 12
10. Forewing veins only slightly inflated (Figs.
234, 240); Holarctic Erebiini
Forewing veins inflated (Fig. 224, for example) 11
MEM. AMER. ENT. SOC, 24
74 THE SATYRIDAE
11. Midtibia twice or more the length of the first
midtarsal subsegment; Holarctic Coenonymphini
Midtibia less than twice as long as the first
midtarsal subsegment; Holarctic Maniolini
12. Forewing with a double-pupilled ocellus in space
Mi-M a or Mi-Ma; Old World 13
Forewing with no ocellus in space Mi-M.i, or if
present only single-pupilled; New World Euptychiini
13. Eyes naked Ypthimini
Eyes hairy genus Palaeonympha
Tribe Hypocystini, new tribe
(Figures 140-169)
The Hypocystini are here considered to be the most primitive
members of the Satyrinae, primarily because of the configurations
of the male and female forelegs. Although these butterflies are not
closely allied to any other tribe within the Satyrinae, they seem to
connect the Ypthimini- and Satyrini-sections. The Xenica-series in
the present tribe connects with the Satyrini-section by pattern and
structural similarities, whereas the Hypocysta-series exhibits an even
closer approach to the Ypthimini-section. The New Guinean hypo-
cystine genus Lamprolenis shows many characteristics in common
with the elymniine tribes Mycalesini and Lethini. This genus has the
typical mycalesine androconial patch along the anterior portion of
the hindwing, and the general facies are reminiscent of the mycalesines
or of Mandarinia. This similarity may offer an explanation of the
evolution of the Satyrinae and will be discussed in greater detail later.
Parallel evolution is shown very well by the New Zealand mem-
bers of this tribe, with the exception of Dodonidia. The other
members, living in montane situations, have evolved into butterflies
very like the Holarctic genus Erebia — indeed, several were placed in
that genus by early authors. Because the New Zealand "coppers" are
placed in the Holarctic genus Lycaena Fabricius (H. K. Clench, per-
sonal communication), the possibility that Argyrophenga and espe-
cially Erebiola and Percnodaimon are indeed referable to Erebia
had to be examined, but there is no morphologic evidence to support
their inclusion in the Holarctic genus. The New Zealand "Erebia"
are highly modified Hypocystini, most nearly related to the Australian
members of the tribe.
LEE D. MILLER
75
Hypocysta
Platypthima
Zipaetis
Hyalodia
Xenica
Heteronympha
Dodonidia
Argynnina
Argyrophenga
Fig. 140. Satyrinae: Hypocystini. Relative lengths of the femur +
tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip-
pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances
the ML value is unity. The top bar for each genus represents the measurements
obtained from males, the bottom bar those from females.
Members of the New Guinean hypocystine fauna (Platypthima,
Erycinidia, Piehdopsis and Lamprolenis) have evolved into bewil-
dering butterflies. In fact, in wing shape and general appearance
Erycinidia is a classic riodinid, and Pieridopsis is a fine pierid
"mimic".
The less miniaturized forelegs in both sexes serves to distinguish
the hypocystines from all other Satyrinae. The forewing cell is usually
deeply excavate in this tribe, a characteristic shared with such
Ypthimini-section tribes as the Ypthimini and Coenonymphini, but
the hindwing crossvein mrraa is produced marginad along M 2 , as
in much of the Satyrini-section. Forewing vein 2A is usually inflated
basad, recalling the Mycalesini, but the inflated part is not sub-
MEM. AMER. ENT. SOC, 24
76
THE SATYRIDAE
quadrate in the hypocystines. A few of the Xenica-series have little
or no inflation of forewing vein 2A.
All members of the Hypocystini are found in the Australian
region, a few being found as far west as Indonesia. Most of the
species are found in New Zealand, Australia, New Guinea and the
associated oceanic islands. Two series of genera are recognized.
Figs. 141-149. Satyrinae: Hypocystini. 141. Hypocysta euphemia West-
wood and Hewitson, $ venation. 142. H. euphemia, antennal club. 143. H.
euphemia, palpus. 144. H. euphemia, S foreleg. 145. H. euphemia, 2 fore-
tarsus. 146. H. euphemia, midleg. 147. Erycinidia gracilis Rothschild and
Jordan, 6 venation. 148. Lamprolenis nitida Godman and Salvin, S vena-
tion. 149. Pieridopsis virgo Rothschild and Jordan, 6 venation.
The Hypocy sta-sches, sensu stricto, most of which are exclusively
tropical insects, are characterized by the generally absent tibial spurs,
LEE D. MILLER
77
the longer third segment of the palpus and hindwing vein Ma arising
nearer M3 than Mi.
The Xenica-series is restricted to the temperate part of the region
and characterized by the development of the tibial spurs, the shorter
third segment of the palpus and (in many genera) hindwing vein
M-, which arises midway between Mi and M3.
The Hypocystini are characterized as follows:
The eyes are generally naked, although they are hairy in such
diverse genera as Argynnina, Heteronympha, Nesoxenica and Pla-
typthima. The antennae are generally two-fifths to half the length
of the forewing costa, but the antennae of Argyronympha are three-
fifths the length of the costa. The antennal club is well developed in
most genera, occupying the distal one-fifth to one-third of the antenna
and expanded to three to four times the width of the shaft (just over
twice the width of the shaft in Lamprolenis) . The club is eccen-
trically developed in some New Zealand genera (Figs. 163, 166,
1 69 ) . The third segment of the palpus is variously one-half to
one-third the length of the second in the Hypocysta-series (Fig. 143)
and one-fourth to one-fifth the length of the second in the Xenica-
series (Fig. 157). The hairs of the second segment of the palpus
vary from less than the width of the segment to more than twice as
long as the segment is wide in the Hypocysta-series to over four times
the width of the segment in some of the New Zealand genera.
Figs. 150-155. Satyrinae: Hypocystini, all Zipaetis saitis Hewitson. 150.
$ venation. 151. antennal club. 152. palpus. 153. $ foreleg. 154. 2
foretarsus. 155. midleg.
MEM. AMER. ENT. SOC, 24
78 THE SATYRIDAE
The male foreleg is moderately well developed (Fig. 140); the
tibia is usually longer than the femur, and the tarsus is usually mono-
merous, but may have as many as four subsegments. The female
foreleg is also well developed (Fig. 140) with a pentamerous, only
slightly clubbed tarsus bearing spines on the first four subsegments
(Fig. 145, for example) — there are more than one pair of spines
on each of the first four subsegments in Dodonidia (Fig. 164). The
midlegs are as long as, or longer than, the hindlegs (Fig. 140). The
midtibia is from less than twice to more than twice the length of the
first midtarsal subsegment, and the tibia is smooth dorsad. Tibial
spurs are generally well developed in the Xenica-series (Fig. 160),
including the New Zealand genera; but the spurs are generally absent
in the Hypocy st a-series (Fig. 146), but they are weakly developed
in Pieridopsis and moderately developed in Zipaetis (Fig. 155). No
spine is developed on the dorsal, distal end of the midtibia.
The forewing cell is excavate, often deeply so, and is from more
than half to more than three-fifths the length of the forewing costa.
In the Hypocysta-series, Argyrophenga and Dodonidia the forewing
radial veins arise in two branches from the cell; in the other genera
the radials arise in three branches. Forewing veins Rs and Mi are
connate in Argyrophenga, Erebiola and Percnodaimon, and separate,
though often approximate, in the other genera. Vein 1VL> arises
nearer, often much nearer, Mi than Ms, and Cui arises a little nearer
M:s than Cun. Vein Sc is characteristically inflated in all genera but
Dodonidia (Fig. 162), and the cubital stem and 2A are inflated in
such diverse genera as Argyrophenga (Fig. 167), Xenica (Fig. 156),
Geitoneura (Fig. 161) and most of the Hypocysta-series (Fig. 141,
for example ) .
The hindwing cell is usually produced at m 2 -m3, but it is more
blunted in Dodonidia (Fig. 162) and Zipaetis (Fig. 150), and the
cell is usually half to three-fifths the length of the wing measured to
the origin and end of M 3 , respectively. The relative lengths of the
hindwing veins Sc + Ri and 3 A are variable. Veins M3 and Cui arise
separately, and M 2 commonly arises midway between Mi and Ms,
but nearer the latter in the New Zealand "Erebia" and the Hypocysta-
Figs. 156-169. Satyrinae: Hypocystini. 156. Xenica achanta (Donovan),
$ venation. 157. X. achanta, palpus. 158. X. achanta, £ foreleg. 159. X.
achanta, 2 foretarsus. 160. X. achanta, midleg. 161. Geitoneura klngii
LEE D. MILLER
79
(Guerin), 8 venation. 162. Dodonidia helmsii Butler, S venation. 163. D.
helmsii, antennal club. 164. D. helmsii, 9 foretarsus. 165. Percnodaimon
pluto (Fereday), $ venation. 166. P. pluto, antennal club. 167. Argy-
rophenga antipodum Doubleday, $ venation. 168. Erebiola buried Fereday,
$ venation. 169. E. butleri, antennal club.
MEM. AMER. ENT. SOC, 24
80 THE SATYRIDAE
series.
The pattern is variable and highly modified. Many of the species
are illustrated by Waterhouse and Lyell (1914) and by Fruhstorfer
(191 2[ 191 2- 191 5]). There is a remarkable resemblance between
Percnodaimon, Erebiola and Argyrophenga and the Holarctic Erebia.
Genera Included in the Hypocystini
Hypocysta-series, sensu stricto
Argyronympha Mathew, 1886: 346. Type-species: Argyronympha pulchra
Mathew, designated by Hemming, 1943: 23.
Erycinidia Rothschild and Jordan, 1905: 457. Type-species: Erycinidia gra-
cilis Rothschild and Jordan, by monotypy.
Harsiesis Fruhstorfer, 1912(1912-1915): 299. Type-species: Hypocysta hy-
geia Hewitson, by monotypy.
Hyalodia Jordan, 1924: 285. Type-species: Hypocysta tenuisquamosa Joicey
and Talbot, by monotypy.
Hypocysta Westwood and Hewitson, 1850, in Doubleday, Westwood and
Hewitson, 1846-1852: pi. 67. Type-species: Hypocysta euphemia
Westwood and Hewitson, by monotypy.
Lamprolenis Godman and Salvin, 1880: 610. Type-species: Lamprolenis
nitida Godman and Salvin, by monotypy.
Pieridopsis Rothschild and Jordan, 1905: 457. Type-species: Pieridopsis virgo
Rothschild and Jordan, by original designation.
Platypthima Rothschild and Jordan, 1905: 458. Type-species: Platypthima
ornata Rothschild and Jordan, by original designation.
Zipaetis Hewitson, 1863(1856-1876): [100]. Type-species: Zipaetis saitis
Hewitson, designated by Butler, 1868a: 194. This genus is exception-
ally aberrant and is only provisionally placed here.
Xenica-series
Argynnina Butler, 1867d: 165. Type-species: Lasiommata hobartia Westwood
and Hewitson, designated by Butler, 1868a: 196.
Argyrophenga Doubleday, 1845: 307. Type-species: Argyrophenga antipodum
Doubleday, by monotypy.
Dodonidia Butler, 1884: 172. Type-species: Dodonidia hehnsii Butler, by
original designation.
Erebiola Fereday, 1879: 128. Type-species: Erebiola butleri Fereday, by
original designation.
Geitoneura Butler, 1867d: 164. Type-species: Satyrus klugii Guerin, desig-
nated by Butler, 1868b: 166.
LEE D. MILLER 81
Heteronympha Wallengren, 1858: 78. Type-species: Papilio merope Fabricius,
designated by Butler, 1868a: 195.
=Hipparchioides Butler, 1867c: 125. Type-species: Papilio merope
Fabricius, designated by Butler. 1868b: 99.
Nesoxenica Waterhouse and Lyell, 1914: 35. Type-species: Lasiommata
leprea Hewitson, by original designation.
= Xeniconympha Noricky, 1923: 60. Type-species: Lasiommata
leprea Hewitson, by original designation.
Oreixenica Waterhouse and Lyell, 1914: 41. Type-species: Lasiommata (?)
lathionella Westwood, by original designation.
Paratisiphone Watkins, 1928: 615. Type-species: Lasiommata lyrnessa Hewit-
son, by original designation.
Percnodaimon Butler, 1876: 152. Type-species: Erebia pluto Fereday, by
original designation.
Tisiphone Hiibner, [ 1 8 1 9] ( 1816-1826) : 60. Type species: Papilio abeona
Donovan, designated by Butler, 1868b: 71.
Xenica Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852:
387. Type-species: Papilio achanta Donovan, designated by Scudder.
1875a: 289.
Tribe Ypthimini, new tribe
(Figures 170-188)
This tribe lies at the base of the Ypthimini-section of the Satyrinae
and seems in some respects to connect with the Hypocysta-sevies of
the last tribe. In its turn the Ypthimini appears to have given rise
to the Holarctic Erebiini and Coenonymphini and the Neotropical
Euptychiini; the Maniolini were evolved later from a coenonymphine
stock.
The Ypthimini are generally characterized by a pattern feature:
the forewing ocellus in space M1-M3 has a double pupil of white or
bluish-white scales. Forewing vein 2A is not inflated in the ypthi-
mines, whereas in the Coenonymphini and many Euptychiini this
vein is greatly distended. While the hindwing crossvein ms-mn is
produced at M2 in the Hypocysta-series, the crossveins are aligned in
this tribe and in other members of the Ypthimini-section, resulting in
a straight cell. The female foreleg is pentamerous in the present
tribe, a situation which is never found in the coenonymphines and
very rare in either the Maniolini or the Erebiini.
MEM. AMER. ENT. SOC, 24
82
THE SATYRIDAE
0.2 XML, FL
Ypthima
Thympia
Xois
Dallacha
Kolasa
Melampias
Pseudonympha
Strabena
Callerebia
Paralasa
Loxeiebia
Fig. 170. Satyrinae: Ypthimini. Relative lengths of the femur + tibia
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling)
and hindlegs (HL, heavy stippling) of selected genera. In all instances the
ML value is unity. The top bar for each genus represents the measurements
obtained from males, the bottom bar those from females.
All ypthimines are found in the Old World, and most genera are
exclusively tropical. Most species fly in open country; few are found
in scrub country, much less in the deep forest. Three major series
are recognized in the Ypthimini, as follows:
The Ypthima-senes, sensu stricto, characterized generally by their
small size, brown coloration and diffuse androconia. This series is
found throughout the Paleotropics.
LEE D. MILLER 83
The Callerebia-series, characterized by large size, brown color-
ation and androconia arranged in rows on the discal portion of the
forewing (Fig. 184). These species are generally distributed in the
temperate mountains of the Middle and Far East.
The Melampias-series, characterized generally by reddish shades
on the upper surface, more elongate wings, small size and either
diffuse androconia or none. These butterflies are found in South
Africa and Madagascar.
The Ypthimini are characterized as follows:
The eyes are naked. The antennae are short, often less than
two-fifths the length of the forewing costa (only one-fourth as long
in Mashuna) and never as long as half the length of the costa (nearly
half in Paralasa and Dallacha). The antennal club is variously
developed: in such genera as Physcaeneura, Periplysia and Dallacha
the club is less than twice as thick as the shaft, whereas in genera like
Mashuna the club is almost four times as thick as the shaft. The
third segment of the palpus is usually from one-fourth to slightly over
one-third the length of the second, but the third segment is more
than half as long as the second segment in Physcaeneura. The hairs
of the second segment of the palpus are very long, usually three to
five times as long as the segment is wide.
The male foreleg is extremely reduced (Fig. 170), with the tarsus
represented in most genera by a small knob on the end of the tibia
(Fig. 173, for example). In such genera as Mashuna (Fig. 177),
Strabena and Pandima, the tarsus is completely fused to the tibia,
but in the Callerebia-series (Fig. 185) the tarsus is moderately well
represented. The forefemur is as long as, or longer than, the fore-
tibia. The female foreleg is also miniaturized, but it retains five
tarsal subsegments, but the tarsus itself is significantly clubbed (Fig.
174, for example). There are spines on the second, third and fourth
subsegments. The midleg is subequal to the hindleg in the Callerebia-
and Melampias-series, but significantly shorter in the Y pthima-series
(Fig. 170). The midtibia is less than twice the length of the first
midtarsal subsegment, is smooth dorsad, without a spine at the dorsal,
distal end and with well to moderately well developed tibial spurs,
except in Physcaenura (Fig. 179).
The forewing cell is usually produced at the origin of Ms and
not excavate (except in Periplysia, Fig. 180); the cell is about half
MEM. AMER. ENT. SOC, 24
84
THE SATYRIDAE
Figs. 171-180. Satyrinae: Ypthimini. 171. Ypthima philomela (Linne),
S venation. 172. Y. philomela, palpus. 173. Y. philomela, 6 foreleg, in-
cluding trochanter and distal portion of coxa. 174. Y. philomela, 9 foretarsus.
175. Y. philomela, midleg. 176. Mashuna mashuna (Trimen), S venation.
177. M. mashuna, $ foreleg, including trochanter and distal portion of coxa.
178. Xois sesara Hewitson, venation of $ hindwing. 179. Physcaemira panda
(Boisduval), midleg. 180. P. panda, ds venation.
as long as the forewing costa. The forewing radial veins arise in one
or two branches from the cell, and veins Rs and Mi are well separated
at their origins. Vein M2 usually arises midway between Mi and M3,
but nearer Mi in the Melampias-series; Cui arises nearer, or much
nearer, Ms than CU2. Forewing vein Sc is always much thickened
at its base, the cubital stem is generally inflated and 2A never is.
LEE D. MILLER 85
The hindwing cell is straight, longest at the origin of Mi and is
about three-fifths the length of the wing measured to the end of Mn.
Hindwing vein Sc + Ri is as long as, and more commonly longer
than, 3A. Veins Ms and Cui arise separately, but often approximate,
and M- arises somewhat nearer Mi than M3, or midway between the
two veins.
As noted in the general description, the pattern is rather constant.
For illustrations of the various species see Moore ( 1893[1 890-1 893])
and van Son ( 1955 ).
Genera Included in the Ypthimini
Ypthima-sevies, sensu stricto
Dallacha Moore, 1893(1890-1893): 94. Type-species: Yphthima (sic.) hya-
griva Moore, by original designation.
Hemadera Moore, 1893(1890-1893): 107. Type-species: Yphthima (sic.)
narasingha Moore, by original designation.
Kolasa Moore, 1893(1890-1893): 82. Type-species: Satyrus chenui Guerin,
by original designation.
Lohana Moore, 1893(1890-1893): 92. Type-species: Yphthima (sic.) inica
Hewitson, by original designation.
Mashuna van Son, (July) 1955: 159. Type-species: Ypthima mashuna Tri-
men, by original designation.
= Ypthimorpha Overlaet, (Dec.) 1955: 23. Type-species: Ypthima
mashuna Trimen, by original designation.
Nadiria Moore, 1893(1890-1893): 85. Type-species: Ypthima bolanica Mar-
shall, by original designation.
Pandima Moore, 1893(1890-1893): 86. Type-species: Satyrus nareda Kollar,
by original designation.
Thympia Moore, 1893(1890-1893): 58. Type-species: Papilio baldus Fabri-
cius, by original designation.
Xois Hewitson, 1865: 282. Type-species: Xois sesara Hewitson, by monotypy.
Ypthima Hiibner, 1818(1818-1825): 17. Type-species: Papilio philomela
Linne, by monotypy. This name is frequently misspelled in the litera-
ture as "Yphthima". Hemming (1964c: 157-158) makes a strong case
for "philomela" of Hiibner being a misidentification of Ypthima hueh-
neri Kirby. Should the International Commission on Zoological No-
menclature rule that this is the case, Y. huebncri will become the type-
species.
Ypthimomorpha van Son, 1955: 158. Type-species: Ypthima itonia Hewit-
son, by original designation.
MEM. AMER. ENT. SOC, 24
86
THE SATYRIDAE
Figs. 181-188. Satyrinae: Ypthimini. 181. Melampias hyperbius (Lin-
ne), $ venation. 182. Cassionympha cassius (Godart), 3 venation. 183. C.
cassius, 9 foretarsus. 184. Callerebia scanda (Kollar), $ venation. 185.
C. scanda, $ foreleg. 186. C. scanda, ? foretarsus. 187. C. scanda, mid-
leg. 188. Boeberia parmenio (Boeber), $ venation.
Melampias-series
Callyphthima Butler, 1880: 335.
by original designation.
Cassionympha van Son, 1955: 96.
original designation.
Coenyropsis van Son, 1958: 6. Type-species: Satyrus natalii Boisduval, by
original designation.
Type-species: Psendonympha wardii Butler,
Type-species: Satyrus cassias Godart, by
LEE D. MILLER 87
Neita van Son, 1955: 101. Type-species: Pseudonym pha neita Wallengren,
by original designation.
Melampias Hubner, [181 9] ( 1816-1826) : 63. Type-species: Papilio hyperbius
Linne. designated by Scudder, 1875a: 214.
Neocoenyra Butler, 1885: 758. Type-species: Neocoenyra duplex Butler, by
monotypy.
Periplysia Gerstacker, 1871: 358. Type-species: Periplysia leda Gerstacker,
by monotypy.
Pseudonym pha Wallengren, 1857: 31. Type-species: Papilio hippia Cramer,
designated by Butler, 1868a: 194.
Physcaeneura Wallengren, 1857: 32. Type-species: Satyrus panda Boisduval,
by monotypy.
Strabena Mabille, 1887: 10, 22. Type-species: Satyrus tamatavae Boisduval,
by original designation.
Stygionympha van Son, 1955: 137. Type-species: Pseudonympha vigilans
Trimen, by original designation.
Callerebia-series
Argestina Riley, 1922: 469. Type-species: Callerebia waltoni Elwes, by origi-
nal designation.
Boeberia Prout, 1901: 233. Type-species: Papilio parmenio Boeber, by origi-
nal designation.
= Erebomorpha Elwes, 1899: 351. Type-species: Papilio parmenio
Boeber, by original designation. Preoccupied by Erebomorpha
Walker, 1860.
Callerebia Butler, 1867e: 217. Type-species: Erebia scanda Kollar, by origi-
nal designation.
Loxerebia Watkins, 1925: 237. Type-species: Callerebia pratorum Oberthur,
by original designation.
Paralasa Moore, 1893(1890-1893): 103. Type-species: Erebia kalinda Moore,
by original designation.
Genus PAL AEON YMPHA Butler, 1871
(Figures 189-195)
Palaeonympha Butler, 1871: 401. Type-species: Palaeonympha opalina But-
ler, by original designation.
The original description of Palaeonympha pointed to its similarity
to the North American Neonympha (Euptychiini). If Palaeonympha
is a member of the Euptychiini it would be the first known Old World
representative of that tribe. It is tempting to place the present genus
there, since it is superficially very similar to the American Megisto
cymela (Cramer), except for the androconial distribution in opalina.
MEM. AMER. ENT. SOC, 24
THE SATYRIDAE
0.4X ML, FL
ML
HL
IxmLi
Y//////A
fmlmmm
?SMmM<M%&m
1
189
i
Palaeonympha
Figs. 189-195. Satyrinae: triba uncertain, Palaeonympha opalina Butler.
189. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diago-
nal lines), midlegs (ML, light stippling) and hind'egs (HL, heavy stippling).
The ML value is unity. The top bar represents the measurements obtained
from the male, the bottom bar those from the female. 190. & venation. 191.
antenna] club. 192. palpus. 193. 6 foreleg. 194. $ foretarsus. 195. midleg.
There are certain similarities betwesn the Euptychiini and the Ypthi-
mini — indeed, they probably arose from the same basic stock — and it
is also tempting to place Palaeonympha in the latter tribe on purely
geographical considerations, but if the present genus is one of the
ypthimines, it is the only one with hairy eyes. Certainly Palaeo-
nympha is either an ypthimine or a euptychiine, or it may represent
an intermediate tribe, but I cannot place the genus in either tribe
with certainty, and I hesitate to erect a monotypic tribe at this time.
Palaeonympha is characterized as follows:
The eyes are slightly hairy. The antennae are just less than
half as long as the forewing costa. The antennal club is moderately
well developed, occupying the distal quarter of the antenna and two
and a half to three times as thick as the shaft. The third segment of
the palpus is long, about two-fifths the length of the second. The
hairs of the second segment of the palpus are three and a half times
as long as the segment is wide.
LEE D. MILLER 89
The male foreleg is greatly reduced (Figs. 189, 193): the femur
is longer than the tibia, and the tarsus is represented only by a bump
at the end of the tibia. The female foreleg is also miniaturized (Figs.
189, 194) with a pentamerous tarsus, the fifth subsegment of which
is greatly reduced, bearing spines on the second and third subseg-
ments. The hindlegs are slightly longer than the midlegs (Fig. 189).
The midtibia is less than twice the length of the first midtarsal sub-
segment, smooth dorsad, with no spine at the dorsal, distal end and
with well developed tibial spurs.
The forewing cell is very slightly excavate between Mi and M-
and about half as long as the forewing costa. The forewing radial
veins arise from the cell in a single branch, and Rs and Mi are
separate at their origins. Vein M- arises slightly nearer Mi than Ma,
and Cui arises nearer Ma than Cu-. Forewing vein Sc and the cubital
stem are both much inflated basad, but 2A is not.
The hindwing cell is somewhat produced at the origin of M3 and
is about three-fifths the length of the wing measured to the end of
Ms. Hindwing vein Sc + Ri is longer than 3 A, and Ma and Cui
are well separated at their origins. Vein M 2 arises nearer Mi than Ma.
The single species is brown above with a double-pupilled ocellus
in Mi-M- of the forewing and single-pupilled ones in Rs-Mi and
Ma-Cui of the hindwing. There is an androconial patch of mealy
scales along the boundary of the cell and extending out the veins
from Mi to 2A (Fig. 190). The under surface is light brown with
discal and extradiscal transverse reddish-brown bands and well de-
veloped ocelli as on the upper surface, plus very small ones in
forewing spaces Ma-Cui through CU--2A and better developed ones
in hindwing spaces M1-M2, Mi>-Ma and CU2-2A.
Tribe Euptychiini, new tribe
(Figures 196-210)
This tribe shows the greatest affinities to the Old World Ypthi-
mini and was probably derived from a common stock with it.
The euptychiines are less closely related to the Maniolini and Coeno-
nymphini.
The Euptychiini are separable from the Ypthimini on the basis
of the slightly longer and more square-cut forewing cell and the
ocellus in forewng space Mi-Ma and differ from the coenonymphines
MEM. AMER. ENT. SOC, 24
90
THE SATYRIDAE
0.4XML
IxML,
Euptychia
Taygetis
Pindis
Paramecera
Oressinoma
Fig. 196. Satyrinae: Euptychiini. Relative lengths of the femur +
tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip-
pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances
the ML value is unity. The top bar for each genus represents the measure-
ments obtained from males, the bottom bar those from females.
in that the latter have an excavate forewing cell and a greater abor-
tion of the female foreleg, as well as a greater distention of forewing
vein 2A.
All of the genera definitely assigned to the euptychiines are found
in the New World, from southern Canada through the Neotropics to
temperate South America. Most are woods or forest dwellers, in
contrast to most members of the Ypthimini-section. The most com-
prehensive review of the tribe to date is included in Forster ( 1964),
but, unfortunately, it is by no means complete. The Euptychiini
are characterized below.
The eyes are variously naked or hairy. The antennae are short,
just under to just over two-fifths the length of the forewing costa.
The antennal club is usually weakly developed and about twice the
thickness of the shaft (about three times as thick in Cyllopsis). The
third segment of the palpus is about a third as long as the second
segment. The hairs of the second segment of the palpus are variable:
in the tropical groups they are about three times as long as the
segment is wide, but in the Nearctic Me gist o cymela (Cramer) the
hairs are almost five times the width of the segment.
The male foreleg is reduced, with a monomerous, unspined tarsus;
the femur and tibia are of about the same length. The female foreleg
LEE D. MILLER
91
Figs. 197-205. Satyrinae: Euptychiini. 197. Euptychia mollina Hiibner,
8 venation. 198. E. mollina, palpus. 199. E. mollina, $ foreleg. 200. E.
mollina, 2 foretarsus. 201. Cissia penelope (Fabricius). 9 foretarsus. 202.
E. mollina, midleg. 203. Megisto cymela (Cramer), midleg. 204. Cyllopsis
hedemanni R. Felder, S venation. 205. Taygetis mermeria (Cramer), S
venation.
is also reduced, with a pentamerous, clubbed tarsus bearing spines
on the second, third and fourth subsegments (also on the first in at
least Neonympha). The midtibia is one and a half to one and three-
quarters times the length of the proximal midtarsal subsegment. The
midtibia is variously smooth dorsad in most genera to slightly spiny
in such genera as Neonympha and spiny in such genera as Cissia.
The tibial spurs are present and well developed in most species, but
MEM. AMER. ENT. SOC, 24
92 THE SATYRIDAE
absent in Euptychia mollina Hiibner, and there is no spine at the
dorsal, distal end of the midtibia. The relative lengths of the legs
of selected Euptychiini are shown in Fig. 196.
The forewing cell is generally square-cut and varies little from
half the length of the forewing costa. The forewing radial veins
arise from the cell in two branches (rarely one branch), and veins
Rs and Mi arise well separate. Vein M2 arises somewhat nearer Mi
than M a , and Cui arises nearer M 3 than CU2. Forewing veins Sc and
the cubital stem are basally inflated in all genera, and 2A is inflated in
such genera as Taygetis and Oressinoma (Figs. 205, 210, respec-
tively). The latter genus displays an unusual form of inflation of
the veins.
The hindwing cell is straight and produced at the origin of M3;
the cell is half to three-fifths the length of the wing measured to the
end of Mm. Hindwing veins Sc + Ri and 3 A are of about the same
length, Ms and Cui arising well separate and M2 arising nearer,
occasionally much nearer, Mi than M3.
The pattern is reasonably constant. Most species are brown
above — they may be white or iridescent purple, blue in part — with
or without ocelli. The under surface is usually brown with trans-
verse darker markings and a more or less complete complement of
ocelli. Oressinoma typhla Westwood is aberrant: white above and
below bordered with blackish brown and with reddish lunules in the
dark border.
Genera Included in the Euptychiini 1
'Names preceded by asterisks (*) are genera proposed too late for de-
tailed study, but from the original descriptions they are judged to be members
of this tribe.
*Archeuptychia Forster, 1964: 80. Type-species: Papilio c'uena Drury, by
original designation.
*Caerul euptychia Forster, 1964: 92. Type-species: Euptychia caerulea But-
ler, by original designation.
*Capronnieria Forster. 1964: 73. Type-species: Euptychia abretia Capron-
nier, by original designation.
*Cepheuptychia Forster, 1964: 96. Type-species: Euptychia cephus Butler,
by original designation.
*Chlor euptychia Forster, 1964: 119. Type-species: Papilio chloris Cramer,,
by original designation.
Cissia Doubleday, 1848b: 33. Type species: Papilio clarissa Cramer (=Pa-
LEE D. MILLER
93
Figs. 206-210. Satyrinae: Euptychiini. 206. Paramecera xicaque (Rea-
kirt), 6 venation. 207. P. xicaque, palpus. 208. P. xicaque, £ foreleg. 209.
P. xicaque, 2 foretarsus. 210. Oressinoma typhla Westwood, 6 venation.
pilio penelope Fabricius), by monotypy.
= Argyreuptychia Forster, 1964: 123. Type-species: Papilio penel-
ope Fabricius, by original designation.
*Coeruleotaygetis Forster, 1964: 73. Type-species: Euptychia periboea God-
man and Salvin, by original designation.
Cyllopsis R. Felder, 1869: 474. Type-species: Cyllopsis hedemanni R. Felder,
by monotypy.
*Erichthodes Forster, 1964: 118. Type-species: Euptychia erichtho Butler,
by original designation.
Euptychia Hlibner. 1818 (1818-1825): 20. Type-species: Euptychia mol-
lina Hubner, designated by Hemming, 1937: 150. Butler (1868a:
194) designated Papilio herse Cramer as the type, based on Euptychia
Hubner, [1819]( 1816-1826) : 54, but this designation was invalidated
when the appropriate portions of the "Zutrage" were found to be older
than those of the "Verzeichniss".
*Euptychoides Forster, 1964: 97. Type species: Euptychia saturnus Butler,
by original designation.
*Godartiana Forster, 1964: 118. Type-species: Satyrus byses Godart, by
original designation.
*Harjesia Forster, 1964: 78. Type-species: Taygetis blanda Moschler, by
original designation.
*Haywardina Forster, 1964: 109. Type-species: Satyrus necys Godart, by
original designation. Preoccupied by Haywardina Aczel, 1952, but no
replacement name is proposed.
*Hermeuptychia Forster, 1964: 87. Type-species: Papilio hermes Fabricius,
by original designation.
MEM. AMER. ENT. SOC, 24
94 THE SATYRIDAE
*Magneuptychia Forster, 1964: 125. Type-species: Popilio libye Linne, by
original designation.
*Megeuptychia Forster, 1964: 122. Type-species: Nymphalis autonoe Cramer,
by original designation.
Megisto Hiibner, [1819]( 1816-1 826) : 54. Type-species: Papilio eurytus Fa-
bricius (=PapiIio cymela Cramer), designated by Butler, 1868b: 14.
*Moneuptychia Forster, 1964: 92. Type-species: Euptychia soter Butler, by
original designation.
Neonympha Hiibner, 1818(1818-1825): 8. Type-species: Papilio areolatus
Smith and Abbot, designated by Hemming, 1937: 150.
Oressinoma Westwood, 1852, in Doubleday, Westwood and Hewitson, 1846-
1852: 371. Type-species: Oressinoma typhla Westwood, by monotypy.
= Ocalis Westwood, 1852, in Doubleday, Westwood and Hewitson,
1846-1852: 371. Type-species: Oressinoma typhla Westwood, by
monotypy. Listed in synonymy of Oressinoma Westwood.
Paramecera Butler, 1868b: 98. Type-species: Neonympha xicaque Reakirt,
by monotypy.
*Parataygetis Forster, 1964: 79. Type-species: Taygetis albinotata Butler, by
original designation.
*Parenptychia Forster, 1964: 83. Type-species: Papilio hesione Sulzer, by
original designation.
* Parypththimoid.es Forster, 1964: 106. Type-species: Neonympha poltys
Prittwitz, by original designation.
*Pharneuptychia Forster, 1964: 91. Type-species: Satyrus phares Godart, by
original designation.
Pindis R. Felder, 1869: 475. Type-species: Pindis squamistriga R. Felder, by
monotypy.
*Posteuptychia Forster, 1964: 137. Type-species: Euptychia mycalesoides
Felder, by original designation.
*Posttaygetis Forster, 1964: 74. Type-species: Papilio penelea Cramer, by
original designation.
*Praefaunula Forster, 1964: 137. Type-species: Euptychia armilla Butler, by
original designation.
*Pseudeuptychia Forster, 1964: 86. Type-species: Euptychia languida Butler,
by original designation.
*Pseudodebis Forster, 1964: 75. Type-species: Papilio valentina Cramer,
by original designation.
*Rareuptychia Forster, 1964: 87. Type-species: Euptychia clio Weymer, by
original designation.
*Satyrotaygetis Forster, 1964: 73. Type-species: Euptychia satyrina Bates,
by original designation.
*Splendeuptychia Forster, 1964: 128. Type-species: Euptychia ashna Hew-
itson, by original designation.
*Taygetina Forster, 1964: 77. Type-species: Taygetis hanghaasi Weymer, by
original designation.
LEE D. MILLER 95
Taygetis Hiibner, [1819]( 1816-1826) : 55. Type-species: Papilio mermeria
Cramer, designated by Westwood, 1852, in Doubleday, Westwood and
Hewitson, 1846-1852: 355. Hemming (in press) accepts the designa-
tion of Papilio virgilia Cramer as the type-species by Butler, 1868a: 194,
but the intent of Westwood to designate mermeria as the type-species
is unmistakable.
*Vareuptychia Forster, 1964: 125. Type-species: Euptychia similis Butler,
by original designation.
*Weymerana Forster, 1964: 108. Type-species: Euptychia viridicans Wey-
mer, by original designation.
*Ypthimoides Forster, 1964: 100. Type-species: Neonympha yphthima Felder,
by original designation.
*Zischkaia Forster, 1964: 116. Type-species: Euptychia jumata Butler, by
original designation.
Tribe Coenonymphini, new tribe
(Figures 211-222)
The Coenonymphini are intimately related to the ypthimines
and probably replace them in the north temperate zone savannas of
both hemispheres. There are some relationships suggested with the
maniolines, the latter group occupying scrub country environments:
probably both evolved from a basic ypthimine stock. There is little
to connect the coenonymphines with the Euptychiini or the Erebiini,
two isolated tribes of the Ypthimini-section.
The excavate forewing cell serves to distinguish most coeno-
nymphines from most ypthimines, and the very long midtibia (over
twice as long as the first midtarsal segment in all genera but Aphan-
topus) is unique in the Ypthimini-section. The female foreleg of the
Ypthimini and the Euptychiini is fully formed, having five tarsal sub-
segments, whereas that of the Coenonymphini is lacking at least one
of the subsegments.
The Coenonymphini are exclusively Holarctic and are grassland
species. Almost all are temperate, few (e. g., Coenonympha mix-
turata Alpheraky) being found so far north as the subarctic. The
Coenonymphini are characterized as follows:
The eyes are naked. The antennae are always shorter than half
the length of the forewing costa, and those of Lyela and Triphysa
are less than two-fifths the length of the forewing. The antennal club
is variable in its development: in Aphantopus the club is only about
MEM. AMER. ENT. SOC, 24
96
THE SATYRIDAE
0.2*ML
Fig. 211. Satynnae: Coenonymphini. Relative lengths of the femur +
tibia + tarsus of the forelegs (FL diagonal lines), midlegs (ML, light stip-
pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances
the ML value is unity. The top bar for each genus represents the measure-
ments obtained from males, the bottom bar those from females.
two and a half times as thick as the shaft, five times as thick in Tri-
physa (Fig. 220) and seven times the thickness of the shaft in Lyela.
The third segment of the palpus is generally half to one-third as long
as the second, but the third segment is less than one-sixth the length
of the second in Triphysa and Lyela. The hairs of the second seg-
ment of the palpus are about four times as long as the segment is
wide.
The male foreleg is weakly developed with a monomerous, un-
spined (weakly spined in Lyela) tarsus. The forefemur and fore-
tibia are of about the same length; in some genera slight variations
may be apparent. The female foreleg is reduced and is of two gen-
eral types: in such genera as Coenonympha there are four tarsal sub-
segments which are spined on at least the third and fourth ones,
whereas in other genera like Chortobius, Lyela and Triphysa there
is but one, unspined tarsal subsegment. With the exception of
Aphantopus, the midtibia is very long, greater than twice as long as
the first midtarsal subsegment. The tibial spurs are present and well
developed in all genera but Triphysa, in which they are totally absent.
The midtibia is smooth dorsad and has no spine at the dorsal, distal
end. The hindlegs are significantly longer than the midlegs (Fig.
211).
The forewing cell is deeply excavate and varies little from half
the length of the forewing costa. The forewing radial veins generally
arise in two branches from the cell, but in three branches in Aphan-
LEE D. MILLER
97
topus; veins Rs and Mi are separate at their origins, although they
may be approximate. Vein M- arises nearer Mi than M :! , and Cui
arises midway between Ms and Cu^, or slightly nearer M3. Fore-
Figs. 212-222. Satyrinae: Coenonymphini. 212. Coenonympha oedippus
(Fabricius). 6 venation. 213. C. oedippus, antennal club. 214. C. oedippus,
palpus. 215. C. oedippus, 6 foreleg. 216. Chortobius pamphilus (Fabricius).
9 foretarsus. 217. C. oedippus, midleg. 218. C. oedippus, 9 foretarsus. 219.
Triphysa phryne (Pallas), 6 venation. 220. T. phryne, antennal club. 221.
T. phryne, midleg. 222. Aphantopus hyperantus (Linne), 6 venation.
wing veins Sc, the cubital stem and 2A are all inflated, though the
latter may be only slightly thickened.
The hindwing cell is square-cut distad and half to three-fifths
the length of the wing to the end of M3. Hindwing vein Sc + Ri is
MEM. AMER. ENT. SOC, 24
98 THE SATYRIDAE
as long as, or longer than, 3A. Veins Ms and Cui are separate,
but close together, at their origins, and M2 arises nearer, or much
nearer, Mi than Ma.
The pattern is fairly constant. The upper surface is brown to
tawny with few, if any, ocelli. The under surface of the forewings
is likewise brown to tawny, and the hindwings are brown to grayish-
olive, often with a light mesial band, with a more or less developed
series of ocelli on both wings. Triphysa phryne (Pallas) is the only
species to show marked sexual dimorphism: the male is brown with
a more complete set of ocelli than is usual in the tribe, and the female
is white with a full complement of ocelli.
Genera Included in the Coenonymphini
Aphantopus Wallengren, 1853: 30. Type-species: Papilio hyperantus (mis-
spelled "hyperanthus") Linne, by monotypy.
Chortobius [Dunning and Pickard], 1858: 5. Type-species: Papilio pamphilus
Linne, designated by Moore, 1893 (1890-1893): 51-52.
Coenonympha Hiibner, [18 19] ( 1816-1826) : 65. Type-species: Coenonympha
oedipe Hiibner (=Papilio oedippus Fabricius), designated by Butler,
1868a: 194 (as Papilio geticus Esper).
Dubierebia Muschamp, 1915b: 20. Type-species: Coenonympha myops
Staudinger, by original designation.
Lyela Swinhoe, 1908: 60. Type-species: Lyela macmahoni Swinhoe, by origi-
nal designation.
Sicca Verity, 1953: 83. Type-species: Papilio dorus Esper, by original desig-
nation. Proposed as a subgenus of Coenonympha Hiibner.
Triphysa Zeller, 1850: 308. Type-species: Papilio tircis Stoll (= Papilio
phryne Pallas), designated by Butler, 1868a: 194.
=Phryne Herrich-Schaffer, 1844 (1843-1856): 90. Type-species:
Papilio tircis Stoll (=Papilio phryne Pallas), by monotypy. Pre-
occupied by Phryne Meigen, 1 800.
Tribe Maniolini Hampson, 1918
(Figures 223-232)
Maniolinae Hampson, 1918: 385; Maniolidi Verity, 1953: 228, 237.
This tribe is most closely related to the Coenonymphini; probably
both tribes evolved in the Holarctic from a basic ypthimine stock.
The maniolines are much more distantly related to either the Eupty-
chiini or the Erebiini.
The maniolines are distinguished from other members of the
LEE D. MILLER
99
Ypthimini-section by forewing vein M-2 arising midway between Mi
and M.i (in other tribes M2 arises nearer Mi), and by the distribu-
tion of androconia. Whereas the Euptychiini and the Ypthimini
0.2 x ML, FL
ML
HL IxML,
1
1
r
v//A
1
1
Maniola
Cercyonis
Fig. 223. Satyrinae: Maniolini. Relative lengths of the femur + tibia
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML
value is unity. The top bar for each genus represents the measurements
obtained from males, the bottom bar those from females.
have a pentamerous female foreleg, this tribe shows leg abortion in
all genera, like the Coenonymphini.
Cercyonis is considered by many authors (e.g., Hemming, in
press ) to be biologically synonymous with Minois Hiibner. The latter
genus, however, is a member of the Satyrini, as shown by the pro-
duction of the hindwing cell along vein Mi> and the typically satyrine
configuration of the midleg (Fig. 301). The midleg of Cercyonis
is typical of the Maniolini (Fig. 228) and has nothing in common
evolutionary with Minois. The proper assignment of Cercyonis
again demonstrates the necessity of looking beyond mere superficial
similarities.
The maniolines are found throughout the north temperate zone,
and a few members are also found in the Arctic. These butterflies
seem to replace the grassland Coenonymphini in scrublands. The
Maniolini are characterized as follows:
The eyes are naked. The antennae are greater than two-fifths,
but less than half, the length of the forewing costa. The antennal club
is weakly developed, from less than twice to about two and a half
times the thickness of the shaft, and occupying the distal third to a
quarter of the antenna. The third segment of the palpus is about
one-third of the length of the second. The hairs of the second seg-
ment of the palpus are long, about four to five times as long as the
segment is thick.
The male foreleg is greatly reduced, the femur and the tibia are
MEM. AMER. ENT. SOC, 24
100
THE SATYRIDAE
of approximately the same length and the tarsus is monomerous and
unspined. The female foreleg is also greatly reduced with a mono-
or dimerous tarsus without spines. The mid- and hindlegs are of
Figs. 224-232. Satyrinae: Maniolini. 224. Maniola jurtina (Linne),
$ venation. 225. M. jurtina, palpus. 226. M. jurtina, $ foreleg. 227. M.
jurtina, 9 foretarsus. 228. M. jurtina, midleg. 229. Pyronia tithonus (Lin-
ne), $ venation. 230. Cercyonis pegala (Fabricius), 6 venation. 231. C.
pegala, S foreleg. 232. C. pegala, 2 foretarsus.
about the same length (Fig. 223). The midtibia is somewhat less
than twice as long as the basal midtarsal subsegment, without spines
dorsad or at the dorsal distal end and with well developed tibial spurs.
The forewing cell is somewhat excavate and rather uniformly half
LEE D. MILLER 101
as long as the forewing costa. The forewing radial veins arise in
three branches from the cell, and veins Rs and Mi are separate at
their origins. Vein M- arises about midway between Mi and M:s.
and Cui arises nearer Ma than Cu-. Forewing vein Sc is always
inflated basally; the cubital stem and 2A may be greatly inflated
(Maniola, Fig. 224) or not noticeably inflated (Cercyonis, Fig. 230).
The hindwing cell is produced at the origin of Ma and varies from
less than half to as long as three-fifths the length of the wing measured
to the end of Ma. Vein Sc + Ri is longer than 3 A, and Ma and Cui
are well separated at their origins. Vein M- arises nearer Mi than Ma.
The upper surface is either tawny with brown margins or brown
with or without a yellow discal patch on the forewing with the ocelli,
if they are present, in the forewing spaces Mi-Ma and Cua-2A and on
the hindwing at the tornus. Beneath the wings are light brown with
darker brown striae and moderately well developed ocelli on both
wings.
Genera Included in the Maniolini
Cercyonis Scudder, 1875b: 241. Type-species: Papilio alope Fabricius ( =
Papilio pegala Fabricius), by original designation.
Hyponephele Muschamp, 1915a: 156. Type-species: Papilio lycaon Rottem-
burg, by monotypy.
I data deLesse. 1952: 72. Type-species: Pyronia cecilia Vallantin. by original
designation. Proposed as a subgenus of Pyronia Hiibner.
Maniola Schrank, 1801: 152. Type-species: Maniola lemur Schrank (=Papilio
jurtina Linne), designated by Scudder. 1875a: 211.
= Epinephele Hiibner, [1819]( 1816-1826) : 59. Type-species: Pa-
pilio janira Linne (= Papilio jurtina Linne), designated by Butler,
1868a: 194. Note that many authors have misspelled this name
as "Epinephile" or "Epinephild".
Pasiphana de Lesse, 1952: 72. Type-species: Hipparchia bathseba Fabricius,
by original designation. Proposed as a subgenus of Pyronia Hiibner.
Pyronia Hiibner, [ 1 8 1 9] < 1816-1826) : 59. Type-species: Pyronia tithone
Hiibner (=Papilio tithonus Linne), designated by Scudder, 1875a: 261.
Tribe Erebiini Tutt, 1896
(Figures 233-246)
Erebiinae Tutt, 1896: 87. 402.
This tribe shows certain affinities with the Ypthimini and to the
Satyrini-section, but is closer to the former; the similarity to the latter
MEM. AMER. ENT. SOC, 24
102
THE SATYRIDAE
is probably convergence. Structurally no group is exceptionally close
to the erebiines, except the Callerebia-series of the Ypthimini.
0.2*ML FL
Erebia
Medusia
Phorcis
Gorgo
Atercoloratus
Fig. 233. Satyrinae: Erebiini. Relative lengths of the femur + tibia
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML. light stippling) and
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML
value is unity. The top bar for each genus represents the measurements
obtained from males, the bottom bar those from females.
The erebiines may be distinguished from other satyrines of the
Ypthimini-section by the extremely short third palpal segment, which
is less than a fourth as long as the second segment in all genera. The
facies of the species are characteristic, but, as indicated in the dis-
cussions of the Dirini, the Hypocystini and the Pronophilini, some
other genera outside the present tribe share the dark coloration,
rounded wings and alpine habits with these butterflies. The very
slight inflation of the forewing veins, even of Sc, is atypical of the
Ypthimini-section.
All of the erebiines are Holarctic, and many species are found
in the high Arctic, as well as in alpine situations; few are at home in
truly temperate climates. Warren (1936) has admirably mono-
graphed this group. The Erebiini are characterized as follows:
The eyes are naked. The antennae are from just over two-fifths
to greater than half the length of the forewing costa. The antennal
club is moderately well to very well developed, generally occupying
the distal quarter of the antenna and inflated as much as five times
the thickness of the shaft. The third segment of the palpus is very
short, from one-quarter to one-sixth the length of the second. The
LEE D. MILLER
103
Figs. 234-246. Satyrinae: Erebiini. 234. Erebia ligea (Linne), 6 vena-
tion. 235. E. ligea, antennal club. 236. E. ligea. palpus. 237. E. ligea, 8
foreleg. 238. E. ligea, 9 foretarsus. 239. E. ligea, midleg. 240. Phorcis
epistygne Hiibner, <5 venation. 241. P. epistygne, S foreleg. 242. P. epi-
stygne, 9 foretarsus. 243. P. epistygne, midleg. 244. Atercoloratus alini
(Bang-Haas), 6 venation. 245. A. alini, palpus. 246. A. alini, $ foreleg, in-
cluding trochanter and distal portion of coxa.
MEM. AMER. ENT. SOC, 24
104 THE SATYRIDAE
hairs on the second segment of the palpus are long, between three
and five times as long as the segment is wide.
The male foreleg is reduced with a mono- or dimerous, unspined
or weakly spined tarsus; the femur is generally as long as, or longer
than, the tibia. The development of the female foreleg is variable,
though the leg itself is always greatly miniaturized (Fig. 233).
There are two to five tarsal subsegments with spines on all but the
last one. The midleg is about as long as, and frequently longer
than, the hindleg (Fig. 233). The midtibia is from just under to
well over one and a half times the length of the first midtarsal sub-
segment. The midtibia never bears a spine at its dorsal, distal end
and is usually smooth dorsad, but slightly spiny in Phorcis (Fig. 243).
The tibial spurs are present and well developed.
The forewing cell is slightly excavate and varies little from half
the length of the forewing costa. The forewing radial veins arise in
three branches from the cell, and veins Rs and Mi are well separated
at their origins. Vein M_> arises much nearer Mi than Ms, and Cui
arises nearer M3 than Cu^>. Forewing vein Sc is always more or less
inflated, the cubital stem is somewhat inflated and 2A hardly at all.
The hindwing cell is produced at the origin of M3 and is half to
three-fifths as long as the wing measured to the end of Ms. Vein
Sc + Ri is longer than 3 A, and M3 and Cui are well separated at
their origins. Vein M2 arises nearer Mi than Ms.
These are dark insects with very little pattern on either surface
except scattered ocelli and a forewing tawny to russet shad 2; either or
both pattern features may be absent. Illustrations of the species are
given by Warren (1936).
Genera Included in the Erebiini
Atercoloratus Bang-Haas, 1938: 178. Type-species: Coenonympha alini Bang-
Haas, by original designation.
Erebia Dalman, 1816: 58. Type-species: Papilio ligea Linne, by original
designation.
= Epigea Hiibner, [1819]( 1816-1826) : 62. Type-species: Papilio
ligea Linne, designated by Hemming, 1933: 198.
Gorgo Hiibner, [18 19] ( 1816-1826 ) : 64. Type-species: Papilio ceto Hiibner,
designated by Hemming. 1933: 198.
Marica Hiibner, [1 8 19] ( 1816-1826) : 63. Type-species: Papilio stygne Ochs-
enheimer, designated by Hemming, 1933: 198.
Medusia Verity, 1953: 179. Type-species: Papilio medus Schiffermiller, by
original designation. Proposed as a subgenus of Erebia Dalman.
LEE D. MILLER 105
Oreina Westwood, 1841: 76. Type-species: Papilio cassiope Fabricius, desig-
nated by Butler, 1868a: 194.
Phorcis Hubner, [ 18 19]( 181 6-1826) : 62. Type-species: Phorcis epistygne
Hiibner, designated by Hemming, 1933: 198.
Simplica Verity, 1953: 194. Type-species: Papilio epiphron Knoch, by origi-
nal designation. Proposed as a subgenus of Erebia Dalman.
Syngea Hubner, [18 1 9] ( 1816-1826) : 62. Type-species: Papilio pronoe Esper,
designated by Hemming, 1933: 198.
Triariia Verity. 1953: 186. Type-species: Papilio triarius de Prunner, by
original designation. Proposed as a subgenus of Erebia Dalman.
Truncaefalcia Verity, 1953: 188. Type-species: Papilio aethiops Esper, by
originial designation. Proposed as a subgenus of Erebia Dalman.
Tribe Dirini, new tribe
(Figures 247-259)
This tribe is not as primitive as the hypocystines, as may be seen
form examination of the forelegs of both sexes, even though it seems
to connect the remainder of the Satyrini-section with the South Afri-
can members of the Lethini — a convergence which is probably
ecological and with little phylogenetic significance. The forelegs of
both sexes are greatly reduced, about as much so as in any tribe;
the dirines are nevertheless primitive members of the Satyrini-section.
There is little to connect the Dirini — or any member of the Satyrini-
section, for that matter — with the Hypocystini except the configura-
tion of the hindwing cell (Figs. 248, 254, 257). In common with
the Satyrini, Pronophilini (usually to a lesser extent) and Melanar-
giini are the greatly reduced forelegs, the spiny midtibiae and the well
developed tibial spurs of the present tribe. Some species have a well
developed dorsal, distal spine at the end of the midtibia (Fig. 259),
a characteristic shared with the Satyrini and indicating a close rela-
tionship between these two tribes.
The Dirini are characterized chiefly by the short forewing cell,
always less than half the length of the wing; in other Satyrinae the
cell is half the length of the wing or greater. The forewing cell is
also rounded and reminiscent of that of the Lethini.
All members of this tribe are restricted to South Africa and are
well illustrated by van Son (1955). The Dirini are characterized
as follows:
The eyes are hairy in all genera. The antennae are approximately
two-fifths the length of the forewing costa. The antennal club is
MEM. AMER. ENT. SOC, 24
106
THE SATYRIDAE
0.2XML, FL
Dira
Tarsocera
Torynesis
Fig. 247. Satyrinae: Dirini. Relative lengths of the femur + tibia +
tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML
value is unity. The top bar for each genus represents the measurements ob-
tained from males, the bottom bar those from females.
abrupt, occupying the distal one-fifth to one-third of the antenna
and three to five times as thick as the shaft (Fig. 249). The third
segment of the palpus is usually about a third the length of the
second, but is half as long in Cassus (Fig. 254) and only one-ninth
as long in Torynesis (Fig. 257). The hairs of the second palpal
segment are two or three times as long as the segment is wide.
The male foreleg is greatly reduced (Fig. 247), particularly as
regards the tibia and tarsus. The femur is always more than twice
as long as the tibia, and the tarsus has one or two unspined sub-
segments (Figs. 251, 258). The female foreleg is also greatly re-
duced, frequently as much so as that of the male, with a mono- or
dimerous, unspined tarsus (Fig. 252). The midleg is as long as,
and occasionally much longer than (Torynesis), the hindleg (Fig.
247). The midtibia is almost twice as long as the first midtarsal
subsegment in some genera such as Dira (Fig. 253), but in Tory-
nesis (Fig. 259) and Dingana the midtibia is only slightly longer
than is the proximal midtarsal subsegment. The midtibia is spiny
dorsad and bears well developed tibial spurs; in most genera there is
no spine at the dorsal, distal end of the midtibia, but the spine is
weakly developed in Dingana. The configuration of the midleg in
Dingana and Torynesis strongly suggests that of the Satyrini.
The forewing cell is generally rounded distad and is greater than
two-fifths, but less than half, the length of the forewing costa. The
forewing radial veins arise from the cell in three branches, and Rs
and Mi are well separated at their origins. Vein M2 arises nearer Mi
than M3, and Cui arises nearer M3 than Cu_>. Only forewing vein
LEE D. MILLER
107
Figs. 248-259. Satyrinae: Dirini. 248. Dira clytus (Linne), S venation.
249. D. clytus, antennal club. 250. D. clytus, palpus. 251. D. clytus, $ fore-
leg. 252. D. clytus, $ foretarsus. 253. D. clytus, midleg. 254. Cassus cassus
(Linne), palpus. 255. C. cassus, S venation. 256. Torynesis mintha (Geyer),
$ venation. 257. T. mintha, palpus. 258. T. mintha, $ foreleg. 259. T.
mintha, midleg.
Sc is inflated basad, and it is only slightly thickened.
The hindwing cell is somewhat produced at the origin of M3
and about half the length of the wing measured to the end of Ms.
Hindwing veins Sc + Ri and 3 A are of about the same length, and
veins Ms and Cui arise separately, though they may be approximate.
Vein Mi» arises much nearer Mi than M3.
MEM. AMER. ENT. SOC, 24
108 THE SATYRIDAE
These butterflies are dark colored insects on the upper surface
with ocelli in forewing spaces M1-M3 and Qi2-2A. On the under
surface the forewing ocelli are repeated and the hind wings are pat-
terned by pale mottling and veinal markings. The general aspect
is that of the Erebiini or the Satyrini. The species are figured by
van Son (1955).
Genera Included in the Dirini
Cassus van Son, 1955: 79. Type-species: Papilio cassus Linne, by original
designation.
Dingana van Son, 1955: 70. Type-species: Leptoneura dingana Trimen, by
original designation.
Dira Hiibner, [ 1 819]( 1816-1 826) : 60. Type-species: Dira clyte Hiibner ( =
Papilio clytus Linne), designated by Scudder, 1875a: 157.
= Leptoneura Wallengren, 1857: 31. Type-species: Papilio clytus
Linne, by monotypy.
Tarsocera Butler, 1899: 903. Type-species: Leptoneura cassina Butler, by
monotypy.
Torynesis Butler, 1899: 903. Type-species: Dira mintha Geyer, by monotypy.
=Mintha van Son, 1955: 76. Type-species: Dira mintha Geyer, by
original designation.
Tribe Pronophilini Clark, 1947
(Figures 260-294)
Pronophilinae Clark, 1947: 149.
This tribe was derived from a basic Satyrini-like stock of Old
World origin but is more primitive than any present-day Satyrini,
particularly as regards the forelegs, hence the pronophilines are placed
before the satyrines in this discussion. In addition to the affinity with
the Satyrini, the Pronophilini share some characteristics (though not
particularly important ones) with the South African Dirini, which
are, in turn, the most primitive tribe within the Satyrini-section. The
pronophilines are only very distantly related to the .A>n/ca-series of
the Hypocystini; hence, any trans- Antarctic distribution of the family
is doubtful.
As in the Hypocystini and Dirini, convergent evolution toward
the Holarctic Erebia is shown by several high-altitude pronophilines
of the Lymanopoda-series. Several members of the Elina-series of.
the present tribe show resemblance to members of the Holarctic
LEE D. MILLER
109
0.4XML, FL
Pronophi la
Daedalma
Eteona
Steroma
Gyrocheilus
Elina
Argyrophorus
Neomaenas
Nelia
Lymanopoda
Homoeonympha
Sarromia
Fig. 260. Satyrinae: Pronophilini. Relative lengths of the femur +
tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip-
pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances
the ML value is unity. The top bar for each genus represents the measure-
ments obtained from males, the bottom bar those from females.
Maniolini; indeed, they were long considered members of Maniola
( — Epinephele ) , but on structural grounds these butterflies are per-
fectly valid pronophilines. Many aberrant patterns have evolved in
the great adaptive radiation of this tribe in the Neotropics: the Chilean
MEM. AME1 FNT. SOC, 24
110 THE SATYRIDAE
species Argyrophorous argenteus Blanchard is silver above, while
Lymanopoda samius Westwood is caerulean blue on the upper surface
and members of Drucina resemble nymphalids or danaids more than
they do other satyrids.
The Pronophilini are the only Neotropical satyrids which have
the hindwing crossvein mi-ni2 produced distad at M 2 and hindwing
vein Sc + Ri completely fused throughout its entire length. The
present tribe can be distinguished from the Satyrini by the longer
midtibia — never less than one and a half times the length of the first
midtarsal subsegment — and the lack of a dorsal, distal midtibial spine.
The more aborted forelegs serve to distinguish these butterflies from
the Hypocystini.
All members of the Pronophilini are Neotropical with the excep-
tion of a single genus (Gyrocheilus) which penetrates the southern-
most Nearctic. Many genera, particularly those of the Elina-series,
are found in the temperate parts of South America. Three groups
are recognized within the pronophilines:
The Pronophila-series, sensu stricto, characterized by the gen-
erally broad wings and rounded margins of the hindwings, except
where modified by tails. The vast majority of these butterflies are
found in tropical forests.
The Elina-series, characterized by broad wings, but the hindwings
are usually produced in the anal region. The forelegs are more
reduced than in the Pronophi la-series. Most of these butterflies are
found in the South Temperate region.
The Lymanopoda-series, characterized by the narrow wings,
rounded hindwing margins and reduced forelegs. Most of these
species are found in the Andean highlands.
The Pronophilini are characterized as follows:
The eyes are usually hairy, but are naked in such diverse genera
as Argyrophorus, Amphidecta, Auca, Spinantenna, Manerebia, Idio-
neurula and Gyrocheilus. The antennae are generally two-fifths to
half the length of the forewing costa. The antennal club is well
developed, occupying the distal one-fifth to one-third of the antenna
and between two and five times the thickness of the shaft. The third
segment of the palpus is long for the Satyrini-section, never less than
one-fourth as long as the second segment and frequently over half
as long as the second in such genera as Lasiophila and Idioneurula.
LEE D. MILLER
11
Figs. 261-272. Satyrinae: Pronophilini. 261. Pronophila thelebe Double-
day and Hewitson. 6 venation. 262. P. thelebe, antennal club. 263. P. the-
lebe, palpus. 264. Drucina leonata Butler, palpus. 265. Amphidecta pignera-
tor Butler, palpus. 266. P. thelebe, 6 foreleg. 267. P. thelebe, 9 foretarsus.
268. P. thelebe, midleg. 269. A. pignerator, midleg. 270. Pedaliodes poesia
(Hewitson), 6 venation. 271. P. poesia, $ foreleg. 272. P. poesia, 2 fore-
tarsus.
The hairs on the second palpal segment are highly variable in their
lengths: in such genera as Eteona and Dioriste the hairs are less
than twice as long as the segment is wide, whereas in other genera,
MEM. AMER. ENT. SOC, 24
112
THE SATYRIDAE
such as Gyrocheilus and Manerebia, the hairs may be as much as five
times the width of the segment.
The male foreleg is moderately well developed in most genera
with one (rarely two) tarsal subsegments that are devoid of spines
Figs. 273-279. Satyrinae: Pronophilini. 273. Catargynnis pholoe (Stau-
dinger), $ venation. 274. C. pholoe, palpus. 275. Coracles enyo Hewitson,
8 venation. 276. Steroma bega Westwood, 6 venation. 277. Daedal ma
dinias Hewitson, <$ venation. 278. D. dinias, palpus. 279. Argyrophorous
argenteus Blanchard. c5 venation.
LEE D. MILLER
113
(except in such genera as Cheimas, Gyrocheilus and Pronophila).
The tibia is usually equal to, or longer than, the femur. Members
of the Lymanopoda-series have greatly reduced forelegs with a mono-
merous, unspined tarsus, and the femur is longer than the tibia. The
female foreleg is usually moderately well developed with a penta-
merus, clubbed tarsus bearing spines on the first four subsegments,
but the foreleg of Gyrocheilus is greatly reduced with two unspined
tarsal subsegments (Fig. 283). The relative lengths of the mid-
and hindlegs are variable (Fig. 260). The mid-tibia is rarely less
than twice as long as the first mid-tarsal subsegment, and it is gen-
erally spiny dorsad, but smooth in such diverse genera as Amphidecta
and Penrosada. The tibial spurs are present and well developed,
and there is no spine at the dorsal, distal end of the midtibia.
The forewing cell is usually excavate and half to three-fifths the
length of the forewing costa. The forewing radial veins arise in
three branches from the cell in most genera, but in such genera as
Calisto, Steroma and Argyrophorus the radials arise in two, or even
a single, stems; veins Rs and Mi arise connately (Idioneurula, etc.)
to widely separate (as Gyrocheilus and Pedaliodes) . Vein M2 usually
Figs. 280-283. Satyrinae: Pronophilini. 280. Proboscis propylea (Hew-
itson), 6 venation. 281. Gyrocheilus patrobas (Hewitson), $ venation. 282.
G. patrobas, palpus. 283. G. patrobas, 2 foretarsus.
MEM. AMER. ENT. SOC, 24
114 THE SATYRIDAE
arises midway between Mi and Ms (but much nearer Mi in the
Lymanopoda-series) , and Cui arises nearer M 3 than Cuo in some
genera, midway between the two veins in some and nearer CU2 in
others. Forewing vein Sc is always inflated, often greatly distended,
and the other stalks may also be — all stalks are inflated in such
genera as Steroma and Spinantenna, for example.
The hindwing cell is produced by the distad displacement of
m2-rri3 and half to over three-fifths the length of the wing measured to
the end of M3. Vein Sc + Ri usually is longer than 3 A, and M3 and
Cui are well separated at their origins. Vein M2 arises nearer, gen-
erally much nearer, M3 than Mi.
The pattern is variable and highly modified, and there is con-
vergence between some of the Lymanopoda-series and the Holarctic
Erebia. Most of the species are figured by Weymer, 1912 (1910-
1912).
Genera Included in the Pronophilini 2
' Several genera have not been seen or were described too recently for
critical inclusion in this revision. From the original descriptions it has, how-
ever, been possible to place them in their taxonomic positions. Such genera
are preceded by an asterisk (*).
Pronophila-series, sensu stricto
*Altopedaliodes Forster, 1964: 148. Type-species: Pronophila tena Hewitson,
by original designation.
Amphidecta Butler, 1867a: 404. Type-species: Amphidecta pignerator But-
ler, by monotypy.
*Antopedaliodes Forster, 1964: 151. Type-species: Pedaliodes antonia Stau-
dinger, by original designation.
Calisto Hiibner, 1816(1806-1838): 16. Type-species: Calisto zangis Hiibner,
designated by Butler, 1868a: 194.
Catargynnis Rober, 1892: 284. Type-species: Daedalma pholoe Staudinger,
designated by Hemming, 1943: 23.
=Pseudomaniola Rober, 1892: 222. Type-species: Daedalma pholoe
Staudinger, by original designation. Preoccupied by Pseudomani-
ola Weymer, 1890.
Cheimas Thieme, 1906: 175. Type-species: Oxeoschistus opalinus Staudinger,
by original designation.
Corades Doubleday, 1848a: 115. Type-species: Corades enyo Hewitson, by
monotypy.
*Corderopedaliodes Forster, 1964: 155. Type-species: Pedaliodes corderoi
Dognin, by original designation.
LEE D. MILLER
115
Figs. 284-294. Satyrinae: Pronophilini. 284. Elina vanessoides Blanch-
ard, $ venation. 285. E. vanessoides, palpus. 286. E. vanessoides, $ fore-
leg. 287. E. vanessoides, 2 foretarsus. 288. E. vanessoides, midleg. 289.
Spinantenna tristis (Guerin), 6 venation. 290. Lymanopoda samius West-
wood, 6 venation. 291. L. samius, antennal club. 292. L. samius, palpus.
293. Idioneurula erebioides (Felder and Felder), $ venation. 294. Manerebia
cyclopina Staudinger, $ venation.
MEM. AMER. ENT. SOC, 24
116 THE SATYRIDAE
Daedalma Hewitson, 1858(1856-1876): [85]. Type-species: Daeda'ma dinias
Hewitson, designated by Butler, 18671: 268.
Dioriste Thieme, 1906: 171. Type-species: Pronophila tauropolis Westwood,
by original designation.
Drucina Butler, 1872: 72. Type-species: Drucina leonata Butler, by monotypy.
Eretris Thieme, 1905: 131. Type-species: Pronophila decorata Felder and
Felder, designated by Hemming, 1943: 24.
Eteona Westwood, 1850, in Doubleday, Westwood and Hewitson, 1846-1852:
254. Type-species: Euterpe tisiphone Boisduval, by monotypy.
Gyrochei'.us Butler, 1867f: 267. Type-species: Pronophila patrohas Hewitson,
by original designation. This name was misspelled "Geirocheihts" by
Holland, 1898: 211.
Lasiophila Felder and Felder, 1859: 325. Type-species: Lasiophi'a cirta Felder
and Felder, designated by Scudder, 1875a: 203.
"Muscopedaliodes Forster, 1964: 153. Type-species: Pedaliodes muscosa
Thieme, by original designation.
Oxeoschistus Butler, 1867f: 268. Type-species: Pronophila puerta Hewitson,
by original designation.
Panarche Thieme, 1906: 228. Type-species: Corades tricordata Hewitson, by
original designation.
*Panyapedaliodes Forster, 1964: 157. Type-species: Pronophila panyasis
Hewitson, by original designation.
*ParapedaIiodes Forster, 1964: 153. Type-species: Pronophila parepa Hew-
itson, by original designation.
Pedaliodes Butler, 1867f: 267. Type-species: Pronophila poesia Hewitson, by
original designation.
*Pherepedaliodes Forster, 1964: 149. Type-species: Pedaliodes pheretiades
Smith and Kirby, by original designation.
*Physcopedaliodes Forster, 1964: 152. Type-species: Pronophila physcoa
Hewitson, by original designation.
Polymastus Thieme, 1906: 138. Type-species: Daedalma doraete Hewitson,
by original designation.
*Praepedaliodes Forster, 1964: 152. Type-species: Pronophila phanias Hew-
itson, by original designation.
*Praepronophi!a Forster, 1964: 182. Type-species: Pedaliodes emma Stau-
dinger, by original designation.
Proboscis Thieme, 1906: 168. Type-species: Pronophila propylea Hewitson, by
original designation.
Pronophila Doubleday and Hewitson, 1849, in Doubleday, Westwood and Hew-
itson, 1846-1852: pi. 60. Type-species: Pronophila thelehe Doubleday
and Hewitson, designated by Butler, 18671: 266.
= Mygona Westwood, 1851, in Doubleday, Westwood and Hewitson.
1846-1852: 357. Type-species: Pronophila thelehe Doubleday.
designated by Hemming, 1939: 133.
Pseudosteroma Weymer, 1912(1910-1912): 241. Type-species: Steroma pro-
LEE D. MILLER 1 17
nophila Felder and Felder, designated by Hemming, 1943: 25.
*Punapedaliodes Forster, 1964: 148. Type-species: Pedaliodes albopunctata
Weymer, by original designation.
Steremnia Thieme, 1905: 137. Type-species: Pedaliodes (?) polyxo Godman
and Salvin, designated by Hemming, 1943: 25.
Steroma Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852:
400. Type-species: Steroma bega Westwood, by monotypy.
*Steromapedaliodes Forster, 1964: 148. Type-species: Pedaliodes albonolaia
Godman, by original designation.
Thiemeia Weymer, 1912(1910-1912): 267. Type-species: Pronophila phoro-
nea Doubleday, designated by Hemming, 1943: 25.
Elina-series
Argyrophorus Blanchard, 1852: 30. Type-species: Argyrophorus argenteus
Blanchard, by monotypy.
Auca Hayward, 1953: 30. Type-species: Satyrus pales Philippi, by original
designation.
Elina Blanchard, 1852: 28. Type-species: Elina vanessoides Blanchard, desig-
nated by Butler, 1868a: 194.
*Haywardella Herrera, 1966: 71. Type-species: Satyrus thione Berg, by origi-
nal designation.
Nelia Hayward, 1953: 42. Type-species: Satyrus nemyroides Blanchard, by
original designation.
Neomaenas Wallengren, 1858: 78. Type-species: Neomaenas servilaea Wal-
lengren, by monotypy.
= Stibomorpha Butler, 1874: 204. Type-species: Stibomorpha deco-
rata Butler (= Neomaenas servilaea Wallengren), by original des-
ignation.
Neosatyrus Wallengren, 1858: 79. Type-species: Neosatyrus ambiorix Wal-
lengren, by monotypy.
Pampasatyrus Hayward, 1953: 28. Type-species: Epinephele gyrtone Berg,
by original designation.
*Pseudomanio!a Weymer, 1890: 107. Type-species: Pseudomaniola euripides
Weymer, designated by Hemming, 1943: 25.
= Neomaniola Hayward, 1949: 156. Type-species: Pseudomaniola
euripides Weymer, by original designation. Proposed to replace
Pseudomaniola Weymer, wrongly believed to be preoccupied.
*Quilaphoethosus Herrera, 1966: 69. Type-species: Satyrus monachus Blan-
chard, by original designation.
Spinantenna Hayward, 1953: 38. Type-species: Satyrus tristis Guerin, by
original designation.
Stuardosatyrus Herrera and Etcheverry, 1965: 74. Type-species: Argyro-
phorus williamsianus Butler, by original designation.
MEM. AMER. ENT. SOC, 24
118 THE SATYRIDAE
Lymanopo da-series
Chillanella Herrera, 1966: 71. Type-species: Faunula stelligera Butler, by
original designation.
Cosmosatyrus Felder and Felder, 1867(1864-1867): 495. Type-species: Cos-
mosatyrus leptoneuroid.es Felder and Felder, designated by Butler,
1868b: 59. Herrera (1965: 70) lists the type of this genus as "Cos-
mosatyrus chiliensis chiliensis C. et R. Felder", but chiliensis is a Guerin
name. In any event, the type was properly designated by Butler.
Etcheverrius Herrera, 1965: 62. Type-species: Satyrus chiliensis Guerin, by
original designation.
Faunula Felder and Felder, 1867(1864-1867): 488. Type-species: Faunula
leucoglene Felder and Felder, by monotypy.
Homoeonympha Felder and Felder, 1867(1864-1867): 487. Type-species:
Homoeonympha pusilla Felder and Felder, by monotypy.
Idioneurula Strand, 1942: 389. Type-species: Idioneura erebioides Felder
and Felder, by original designation.
= ldioneura Felder and Felder, 1867(1864-1867): 474. Type-species:
Idioneura erebioides Felder and Felder, by monotypy. Preoccu-
pied by Idioneura Selys, 1860.
Lymanopoda Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-
1852: 401. Type-species: Lymanopoda samius Westwood, designated
by Butler, 1868a: 196.
Manerebia Staudinger, 1897: 139. Type-species: Manerebia cyclopina Stau-
dinger, designated by Hemming, 1943: 24.
Palmaris Herrera, 1965: 67. Type-species: Hipparchia monticolens Butler, by
original designation.
Penrosada Brown, 1944: 255. Type-species: Lymanopoda leaena Hewitson,
by original designation.
*Sabatoga Staudinger, 1897: 143. Type-species: Sabatoga mirabilis Stau-
dinger, by monotypy.
Sarromia Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-
1852: 402. Type-species: Sarromia obsoleta Westwood, by monotypy.
Stygnolepis Strand, 1942: 389. Type-species: Sty gnus humilis Felder and
Felder, by original designation.
= Stygnus Felder and Felder, 1867(1864-1867): 487. Type-species:
Stygnus humilis Felder and Felder, by monotypy. Preoccupied by
Stygnus Perty, 1833.
Tetraphlebia Felder and Felder, 1867 (1864-1867): 489. Typ;-species: Tetra-
phlebia germainii Felder and Felder, by monotypy.
Zabirnia Hewitson, 1877: 92. Type-species: Zabirnia zigomala Hewitson ( =
Lymanopoda acraeida Butler), by monotypy.
= Trophonina Rober, 1892: 222. Type-species: Lymanopoda acrae-
ida Butler, by original designation.
LEE D. MILLER
119
Tribe Satyrini Boisduval, 1836
(Figures 295-308)
Satyrides Boisduval. 1836: 166.
= Hipparchiadae Kirby, 1837: 297.
This tribe is at the center of its section, showing relationships
with the Dirini by virtue of the general facies and the presence of
the heavy spine at the dorsal, distal end of the midtibia, and with
the Pronophilini and much of the Hypocystini in the shape of the
hindwing cell. Through Davidina there is a progressive gradation
from typical Satyrini to the Melanargiini.
0.2xML, FL
Satyrus
Hipparchia
Neominois
Mmois
Karanasa
Davidina
Oeneis
Fig. 295. Satyrinae: Satyrini. Relative lengths of the femur + tibia +
tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML
value is unity. The top bar for each genus represents the measurements ob-
tained from males, the bottom bar those from females.
The weakly developed forelegs' of both sexes serve to distinguish
this tribe from the Hypocystini and much of the Pronophilini, and
the production of the hindwing crossvein m-m a at M- will distinguish
the satyrines from Melanargiini, Dirini and Erebiini. The most char-
MEM. AMER. ENT. SOC, 24
120 THE SATYRIDAE
acteristic features, however, of the Satyrini are on the midleg: all
genera except Chionobas and Oeneis have a heavy spine at the dorsal,
distal end of the midtibia, and in all genera but the O^nm-series and
Davidina the midtibia is less than one and one-fourth times as long
as (and often shorter than) the first midtarsal subsegment.
All members of the Satyrini are Holarctic. The Oeneis-series
is found throughout the Holarctic, but the Satyrus-series is almost
exclusively Palearctic: only Neominois is known from North America.
There is a very interesting pattern similarity between the Euro-
pean genus Minois and the North American Cercyonis, but while
the European butterflies belong to the present tribe, Cercyonis is a
member of the Maniolini. Several authors (e.g., Hemming, in press)
have considered Cercyonis synonymous with Minois, but structural
examination of the two genera does not confirm this conclusion, as
noted in the discussion of Maniolini.
Davidina is intriguing. Several authors have claimed it was a
pierid, but examination of the tarsal claws shows Davidina to be a
satyrid. Pierid tarsal claws are characteristically bifid (see Schatz
and Rober. 1892), whereas those of Davidina are simple, as in most
other satyrids.
Two series are recognized in the Satyrini, as follows:
The Satyrus-series, sensu stricto, distinguished by the extremely
short midtibia and the presence of a heavy spine on the dorsal, distal
end of the midtibia.
The 0<?/7<?/s-series, distinguished by the longer midtibia lacking
the spine at the dorsal, distal end.
The Satyrini are characterized as follows:
The eyes are naked. The antennae are more than two-fifths, but
less than half, the length of the forewing costa. The antennal club
is well developed, occupying the distal one-fourth to one-fifth of the
antenna and three to more than five times as thick as the shaft (Fig.
297). The third segment of the palpus is short, generally one-fourth
to one-fifth as long as the second segment. The hairs of the second
palpal segment are long, two and a half to five times as long as the
segment is wide.
The male foreleg is greatly reduced, the tibia and femur are of
about the same length and the tarsus is mono- to trimerous without
spines (Figs. 299, 306). The female foreleg is also greatly reduced,
as much so as that of the male in several genera (Fig. 295), with
LEE D. MILLER
121
301
Figs. 296-303. Satyrinae: Satyrini. 296. Satyrus actaea (Linne), S
venation. 297. S. actaea, antennal club. 298. S. actaea, palpus. 299. S.
actaea, 6 foreleg. 300. S. actaea, 2 foretarsus. 301. S. actaea, midleg. 302.
Brintesia circe (Fabricius), 6 venation. 303. Neominois ridingsii (Edwards),
5 venation.
the tarsus unspined, or only weakly spined, and comprised of one to
four subsegments. The hindleg is significantly longer than the mid-
leg (Fig. 295). In most genera the midtibia is less than one and
one-fourth times the length of the proximal midtarsal subsegment,
but in Davidina and the Oeneis-szrizs the midtibia is at least half
again as long as the first midtarsal subsegment. The midtibia is very
spiny dorsad, the spines being particularly heavy in this tribe, and
the tibial spurs are present and well developed. There is a charac-
teristic heavy spine at the dorsal, distal end of the midtibia in the
MEM. AMER. ENT. SOC, 24
122
THE SATYRIDAE
Satyrus-series (Fig. 301), which is absent in the Oeneis-series.
The forewing cell is square-cut to slightly excavate and half to
almost three-fifths the length of the forewing costa. The forewing
Figs. 304-308. Satyrinae: Satyrini, all Oeneis noma (Thunberg). 304.
$ venation. 305. palpus. 306. S foreleg. 307. 2 foretarsus. 308. midleg.
radial veins arise from the cell in three branches, and veins Rs and
Mi arise separate, but they may be approximate. Vein M 2 arises
nearer Mi than M3, and Cui arises midway between M :) and CU2,
or slightly nearer the latter. Forewing vein Sc is inflated basad, but
the other veins are little, if at all, thickened.
The hindwing cell is produced by the distad displacement of
ni2-m 3 and half to three-fifths (slightly more in Oeneis) the length
of the wing measured to the end of M3. Veins Sc + Ri and 3 A are
of about the same length, and M :i and Cui are well separated at their
origins. Vein M2 arises nearer Ms than Mi.
The pattern is relatively uniform. The upper surface is shaded
black, brown and/or orange and white with forewing ocelli in spaces
M1-M3 and Cu 2 -2A. The under surface pattern repeats the forewing
ocelli, and the hindwings are cryptically marked. Davidina is white
or off-white with dark brown to black veinal and interveinal lines.
Illustrations of many of the species are given by Moore, 1893(1890-
1893).
LEE D. MILLER 123
Genera Included in the Satyrini
Satyrus-series, sensu stricto
Arethusana de Lesse, 1951: 40. Type-species: Papilio arethusa Esper, by
original designation.
Aulocera Butler, 1867b: 121. Type-species: Satyrus brahminus Blanchard,
designated by Butler, 1868a: 194.
Berberia de Lesse, 1951: 41. Type-species: Satyrus abdelkader Pierret, by
original designation.
Brintesia Fruhstorfer, 1912(1912-1915): 307. Type-species: Papilio circe
Fabricius, by original designation.
Chazara Moore, 1893(1890-1893): 21. Type-species: Papilio briseis Linne,
by original designation.
Davidina Oberthiir, 1879: 19. Type-species: Davidina armandi Oberthur, by
monotypy.
Eumenis Hiibner, [1819]( 1816-1826) : 58. Type-species: Papilio autonoe
Esper, designated by Grote, 1873: 62.
Hipparchia Fabricius, 1807: 281. Type-species: Papilio fagi Scopoli, desig-
nated by Butler, 1868a: 194.
=Nytha Billberg, 1820: 77. Type-species: Papilio hermione Linne
(=Papilio fagi Scopoli), designated by Scudder, 1875a: 231.
=Melcmia Sodoffsky, 1837: 81. Type-species: Papilio fagi Scopoli,
by original designation. Preoccupied by Melania Lamarck, 1799.
Proposed to replace Hipparchia Fabricius, wrongly believed to be
preoccupied.
Kanetisa Moore, 1893(1890-1893): 42. Type-species: Hipparchia digna
Marshall, by original designation.
Karanasa Moore, 1893(1890-1893): 38. Type-species: Satyrus huebneri Feld-
er, by original designation.
Minois Hiibner, [18 19] ( 1816-1826) : 57. Type-species: Papilio phaedra
Linne (=Papilio dryas Scopoli), designated by Butler, 1868a: 194.
Neohipparchia de Lesse, 1951: 40. Type-species: Papilio statilinus Hufnagel,
by original designation.
Neominois Scudder, 1875b: 241. Type-species: Satyrus ridingsii Edwards, by
original designation.
Paroeneis Moore, 1893(1890-1893): 36. Type-species: Chionobas pumilus
Felder, by original designation.
Philareta Moore, 1893(1890-1893): 23. Type-species: Papilio hanifa Nord-
mann (=Papilio anthe Ochsenheimer), by original designation.
Pseudochazara de Lesse, 1951: 42. Type-species: Satyrus pelopea Klug, by
original designation.
Pseudotergumia Agenjo, 1947: unnumbered page at end. Type-species: Pa-
pilio phidia Linne, by original designation.
Satyrus Latreille, 1810: 355. Type-species: Papilio actaea Linne, designated
MEM. AMER. ENT. SOC, 24
124
THE SATYRIDAE
by the action of the International Commission on Zoological Nomen-
clature, 1943. Opinion 142.
Oeneis-series
Chionobas Boisduval, [1833]( 1832-1843) : 182. Type-species: Papilio aello
Hiibner (=Papi!io glacialis Moll), designated by Blanchard, 1840: 457.
Oeneis Hiibner, [ 1819] ( 1816-1 826) : 58. Type-species: Papilio noma Thun-
berg, designated by Butler, 1868a- 196.
Tribe Melanargiini Verity, 1920
(Figures 309-314)
Melanargiinae Verity, 1920: 56.
= Agapetinae Verity, 1953: 3, 46.
This tribe contains some of the most distinctive of all satyrids,
the pattern being unmistakable. In the shape of the hindwing cell,
morphology of the legs and general facies (excluding the white ground
0.2 x ML, FL
Melanargia
Ledargia
Argef ormia
Fig. 309. Satyrinae: Melanargiini. Relative lengths of the femur +
tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip-
pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances
the ML value is unity. The top bar for each genus represents the measure-
ments obtained from males, the bottom bar those from females.
color), these butterflies closely resemble the Dirini and probably
developed from a common stock with them. The spiny legs, reduc-
tion of the forelegs and other structures suggest a strong affinity of
the melanargiines and the Satyrini. There is little to relate this tribe
closely either to the Hypocystini or to the pronophilines.
The pattern of these butterflies sets them apart from all relatives
— white with black marbling and well developed sets of ocelli on
both wings, particularly the hindwings.
All members of the Melanargiini are found in the Palearctic from
LEE D. MILLER
.25
Europe to China and Manchuria. There are more species in Asia
than in Europe. The Melanargiini are characterized as follows:
The eyes are naked. The antennae vary little from half the
length of the forewing costa. The antennal club is rather weakly
developed, less than two to less than three times the thickness of
the shaft. The third segment of the palpus is one-third to one-fourth
the length of the second segment. The hairs of the second segment
of the palpus are about four times as long as the segment is wide.
The male foreleg is greatly reduced, with a single, unspined tarsal
subsegment; the femur is longer than the tibia (Fig. 312). The
female foreleg is as reduced as that of the male with a monomerous,
unspined tarsus (Fig. 313). The relative reduction of the fore-, mid-
and hindlegs is shown in Fig. 309. The midlegs are usually shorter
than the hindlegs (Fig. 309). The midtibia ranges from just under
to just over one and a half times as long as the proximal midtarsal
Figs. 310-314. Satyrinae: Melanargiini, all Melanargia galathea (Linne).
310. 6 venation. 311. palpus. 312. 6 foreleg. 313. 2 foretarsus. 314.
midleg.
subsegment and is very spiny dorsad. The tibial spurs are present
and well developed; there is no heavy spine at the dorsal, distal end
of the midtibia.
The forewing cell is square-cut and approximately half the length
of the forewing costa. The forewing radial veins arise in three
branches from the cell, and veins Rs and Mi are separate, but may
MEM. AMER. ENT. SOC, 24
126 THE SATYRIDAE
be approximate, at their origins. Vein M 2 arises much nearer Mi
than M3, and Cui arises nearer M3 than CU2. If any forewing veins
are inflated, only Sc would be, and that not extensively.
The hindwing cell is produced at the origin of M3 and half to
three-fifths the length of the wing measured to the end of Ms. Veins
Sc + Ri and 3 A are of about the same length, and M3 and Cui are
well separated at their origins. Vein M 2 arises slightly nearer Mi
than M3.
The pattern is quite distinctive: all species are white or cream-
colored marbled with black and with the ocelli well developed. Many
genera have been named, but it seems that one genus would suffice
biologically for all the species.
Genera Included in the Melanargiini
Argeformia Verity, 1953: 47. Type-species: Papilio arge Sulzer, by original
designation. Proposed as a subgenus of Agapetes Billberg (=Mela-
nargia Meigen).
Epimede Houlbert, 1922: 132, 142, 160. Type-species: Arge halimede Mene-
tries, designated by Hemming, 1934a: 143.
Halimede Oberthur and Houlbert, 1922: 192. Type-species: Halimede asiatica
Oberthiir and Houlbert, by monotypy.
Lachesis Oberthur and Houlbert, 1922: 192. Type-species: Lachesis rusci-
nonensis Oberthur and Houlbert ( =Arge lachesis Hiibner), by original
designation.
Ledargia Houlbert, 1922: 157. Type-species: Melanargia leda Leech, by
original designation.
Melanargia Meigen, 1829: 97. Type-species: Papilio galathea Linne, desig-
nated by Kirby, 1894: 41.
=Arge Hiibner, [1819]( 1816-1826) : 60. Type-species: Papilio
psyche Hiibner (= Papilio occitanica Esper), designated by Butler,
1868a: 196. Preoccupied by Arge Schrank, 1802. This name
was suppressed in favor of Melanargia Meigen by the International
Commission on Zoological Nomenclature, 1956, Opinion 400.
= Agapetes Billberg, 1820: 78. Type-species: Papilio galathea Linne,
designated by Scudder, 1875a: 104. This name was suppressed in
favor of Melanargia Meigen by the International Commission on
Zoological Nomenclature, 1956, Opinion 400.
Parce Oberthur and Houlbert, 1922: 193. Type-species: Parce fergana Ober-
thiir and Houlbert (=Melanargia parce Staudinger), by original des-
ignation.
LEE D. MILLER
127
Genera of Uncertain Position
Genus Pamperis Heimlich, 1959
(Fig. 315)
Pamperis Heimlich, 1959: 177. Type-species: Pamperis poaoeneis Heimlich,
by original designation.
Fig. 315. Venation of Pamperis poaoeneis Heimlich (after Heimlich,
1959). Note the aberrant venation pattern discussed in the text.
Having seen no specimens of the type-species of this Neotropical
monotypic genus, I can only evaluate its position in terms of the
original description, which leaves many questions unanswered. Heim-
lich's photographs of the types suggest he was correct, at least
superficially, in placing this genus near Cosmosatyrus Felder and
Felder, and as such, it certainly belongs in the Lymanopoda-seriQs
of the satyrine tribe Pronophilini. However, the venational drawing
given represents either an extraordinary aberration — the veins in the
apical portion of the forewing are arranged like no known satyrid,
and the hindwing lacks a major vein (Fig. 315) — or represent draft-
MEM. AMER. ENT. SOC, 24
128 THE SATYRIDAE
ing errors. Until these structures, as well as others not considered
in the original description, have been studied, it will be quite im-
possible to assign this genus to its proper position among the Satyridae.
Genus Setodocis Billberg, 1820
Setodocis Billberg, 1820: 78. Type-species: Papiiio periboea Fabricius, de-
signated by Scudder, 1875a: 268.
I have no idea to what this genus refers, and neither does anyone
else; periboea is a "lost" species. The best opinion is that of Hem-
ming (in press), "Include at the end of the subfamily Mycalesinae as
a genus dubium." The name is in the literature, so it must be men-
tioned here, but it would seem advisable to suppress it.
The Evolution and Zoogeography of the Satyridae
The twin subjects of evolution and biogeography have provided
biologists since the days of Charles Darwin and Alfred Russel Wal-
lace with the raw material for sundry mental gymnastics and manipu-
lations of the earth's flora, fauna, history and even crust through
space and time. Such exercises, while based on the best available
evidence, are limited by that evidence, and what are proposed as facts
must be considered in actuality only as more or less educated guesses.
The preceding statement is particularly true when one is dealing with
such insects as butterflies, the fossil records of which are almost non-
existent. In drawing evolutionary, and especially zoogeographic, in-
ferences it is necessary to guess the past history of butterflies by com-
parison with fossil histories of vertebrates and to interpret tendencies,
such as the simplification of structures through phylogenetic lines, by
the interpretations given such trends in groups with reasonably docu-
mented fossil records.
Zoogeography, in particular, has been a constant source of
controversy, with several opposing schools of thought attempting to
analyze distributional data in accordance with conflicting theories.
At present there are two major camps of biogeographers which may be
called the Wegener and the Matthew schools, although each theory
has been refined far beyond its originator's proposals.
"Nothing in zoogeography has brought forth more argument or
more demands for an open mind — the other man's mind — than the
idea of continental drift." (Darlington, 1957: 606). This theory,
LEE D. MILLER 129
first put forth by Wegener, states briefly that the continents were a
single landmass which broke apart, and the pieces drifted across the
relatively plastic mantle, chiefly during the Mesozoic and into the
Tertiary, to their present positions. By rotating the present-day
continents and fitting them together it is possible to arrive at a con-
vincingly completed "jig-saw puzzle"; indeed, it was this sort of
piecing together of the landmasses of the world which provided the
basis for the theory in the first place. Caster (1952) summarized
the stratigraphic evidence supporting connections between South
America and Africa, particularly during the Mesozoic, and further
cited the similarity between the shallow water invertebrates of the
two continents, but Dunbar ( 1952: 154) demonstrated by the simi-
larity of shallow water molluscs of Kwajalein, Bikini and Guam that
many shallow water invertebrates can cross deep water "barriers",
probably as free-swimming larvae. Darlington (1957: 607-608)
showed another of the problems with the acceptance of the Wegener
hypothesis as an explanation of the present-day animal distributions,
when he stated, ". . . if a group of animals now occurs only in
Africa and Brazil, it is said to date from and be evidence of a hypo-
thetical African-Brazilian continent. This method depends on an
assumption which Wegenerians usually do not put into words: that
animal distributions are more permanent than land, that animals
move less than continents." This argument more or less summarizes
my own objections to continental drift as an explanation for the dis-
tributions of present-day organisms.
Most American zoogeographers today subscribe to the general
theory of animal dispersal proposed by Matthew (1915) and elab-
orated upon by Hesse, Allee and Schmidt (1951) and Darlington
(1957). Matthew set forth his thesis in the beginning of his paper
(1915: 173).
While the details of Matthew's work have been refined, the
general skeleton has remained intact. Geologic evidence, such as that
presented by Schuchert (1935) for the history of the Neotropics,
has strengthened the Matthew theory. Biogeographers have been
better able to devote their attention to the more knotty problems of
how, when and why organisms arrived where they are.
Since Schuchert (1935) showed the existence of several Ter-
tiary "water gaps" between North and South America and roughly
MEM. AMER. ENT. SOC, 24
130 THE SATYRIDAE
dated them, many authors have considered it an absolute necessity
to close these gaps physically so that organisms might cross them.
This practice, however, has recently come under scrutiny from sev-
eral authors, notably Simpson (1952), who demonstrated mathe-
matically that given a length of time of sufficient duration and a
sufficiently large "feeder" population even the most improbable dis-
persal of a terrestrial organism over water could be a virtual certainty.
While this mathematical exposition does not serve as evidence of a
population of an area via waif dispersal, it shows a method by which
such dispersal could be accomplished. It is of great importance to
the understanding of the aerial dispersal of such insects as butterflies
because it shows that landmasses need never be physically connected
to exchange faunal elements across barriers. Such studies indicate
that reappraisals are in order concerning the necessity of erecting
land bridges across ocean deeps. Certainly physical barriers work
less hardship on such strong flying butterfly genera as Danaus (Da-
naidae), Precis and Vanessa (Nymphalidae), Libythea (Libytheidae)
and Leptotes and Lampides (Lycaenidae) than on the more sedentary
Satyridae and Ithomiidae.
I wish here to call attention to the second paragraph of Matthew's
thesis (1915: 173). Nearly all the principal routes of migration pass
through either the eastern Palearctic or the Indo-Malayan region.
If a group is in a rather generalized condition at the time of its
expansion, it seems illogical that such a group would not leave some
more or less primitive members along its primary path of dispersal.
In the case of biotic interchange between the Palearctic and the
Nearctic, the Paleotropics and the Neotropics, Asia and Australia,
Africa and Australia, these waves of dispersal would have extended
through eastern Asia. Eastern Asia also provides the complete spec-
trum of biotopes, from Arctic tundra to tropical rainforest to the
most arid desert — a prospective "home" for almost any group. No
other region on earth is fed by so many major routes of dispersal,
hence, no other region should be expected to have organisms derived
from so many different sources. This diversity has led many students
(e.g., Darlington, 1957) to consider eastern Asia the major center
of dispersal, a veritable "Garden of Eden". In the following pages
I shall attempt to show that at least the Satyridae need not have
arisen in this region, although eastern Asia certainly played an
LEE D. MILLER 131
important role in the subsequent evolution of the family.
Origin of the Satyridae
Consideration of the origin and subsequent evolution of the
Satyridae must be prefaced by a discussion of their relatives. Two
nymphaloid families, the Morphidae (including the Amathusiidae)
and Ithomiidae, are closely allied to the family under consideration
here and have been characterized at the beginning of the systematic
revision. The evolution of all three families must be considered
together to lay the groundwork for that which follows.
The living nymphaloid families divide basically into two major
groups on larval foodplant requirements. With very few exceptions
(Ehrlich and Raven, 1965) the "true" nymphalids feed on dicoty-
ledonous plants, whereas the larvae of the satyrids utilize monocoty-
ledons exclusively.
Forbes (1932) has assumed that the modern lepidopterous fami-
lies had already differentiated by the Jurassic, and Carpenter (1930)
felt that the Lepidoptera as an order could be traced only as far
back as the Jurassic. Present evidence (Just, 1948) indicates that
the proliferation and differentiation of the angiosperms took place
in the late Mesozoic. The differentiation of the nymphaloid stock
must have been somewhat synchronous with this proliferation of the
flowering plants, but there is some question about this point. Fox (in
lift.) feels that there is no reason to assume that morphological
differentiation may not have preceded plant evolution, i.e., present
monocotyledon feeders may have begun as dicotyledon feeders, or
vice versa. Whether plant or butterfly evolution came first becomes
a moot point, but the potential new niche provided by the evolution
of the two basic stocks of plants must have provided some impetus
for the subsequent, if not the initial, evolution of the butterflies. In
any event, the butterflies are a relatively recent group compared with
the Orthoptera and the Odonata, which were well developed by the
Carboniferous.
The Ithomiidae feed as larvae almost exclusively on the So-
lanaceae (Fox, 1956: 13) and belong on this basis to the "true"
nymphalids, although they are not far removed from the satyrid
complex. These butterflies have reached their highest development
MEM. AMER. ENT. SOC, 24
132 THE SATYRIDAE
in the Neotropics, although a single tribe, the Tellervini, is found in
Australia and New Guinea.
The Satyridae, sensu stricto, overwhelmingly feed as larvae on
Gramineae and Cyperaceae, and all known satyrids feed on mono-
cotyledons. Some members of the family feed on Palmaceae (Elym-
nias and relatives). All known brassolines feed on monocots,
generally utilizing the primitive ones, but a few also feed on bamboo;
none are known to use dicots. The brassolines are exclusively Neo-
tropical, none living farther north than Mexico. The foodplant
requirements of the other primitive Neotropical satyrids (Haeterinae
and Biinae) are unknown, but since they are restricted to the deep
forest they probably do not use grasses, other than possibly bamboo.
There are many other monocotyledons in the rainforest, and these
are probably the food plants.
Structurally, the Indo-Australian Amathusia, Faunis, Taenaris,
etc. belong with the American Morpho in the family Morphidae. All
the known Indo-Australian Morphidae feed on monocotyledons as
larvae, many using Gramineae. In the New World, however, the
morphids (represented only by the nominate genus) predominantly
feed on dicotyledonous plants of the family Canellaceae, Myrtaceae,
Menispermiaceae, Rhamnaceae, Sapindaceae and Leguminoseae.
One species, however, Morpho aega (Fabricius), feeds on bamboo
(Gramineae). Morpho is one of the few nymphaloid genera known
which utilizes both monocotyledons and dicotyledons as larval food;
indeed, the Morphidae is the only family commonly showing such
habits. It seems evolutionary significant that the genus Morpho is
limited to the New World tropics. Furthermore, excluding the
brightly colored and highly modified upper surface of these butter-
flies, the morphids seem to typify the "primitive nymphalid pattern"
of Schwanwitsch (1924): most of the elements of the pattern are
present in the primitive species.
In summary, many of the most primitive members of various
nymphaloid families occur only in the Neotropics. The Neotropical
tribe Tithoreini of the Ithomiidae is as primitive as the Indo-
Australian Tellervini, and the two might well be combined were it
not for their geographic isolation and a few morphologic characters
(Fox, 1956: 20). As shown above, the Neotropical genus Morpho
is the most primitive of the morphids. The most primitive satyrids
are the Neotropical Haeterinae, which preserve the forewing vein 3A.
LEE D. MILLER
133
Fig. 316. The early distributional history of the Haeterinae, Brassolinae
and Biinae in the New World. All movement illustrated in this figure was
during the Cretaceous and earliest Tertiary (open arrows). In succeeding
figures stippled arrows indicate mid-Tertiary movement, whereas solid black
arrows indicate late Tertiary and Quaternary migration. Major areas of the
world are indicated on this map for reference, and no attempt is made to
present exact continental outlines at any given period of geologic time.
The next most primitive groups of satyrids are the biines and the
brassolines and the majority of these, including the most primitive
ones, are New World species. In addition to these satyrids, the
most primitive danaids (Clothilda and the Lycoreinae) are found in
Central America and the Antilles. The remaining danaids are de-
rived from Paleotropical groups which arrived in the Neotropics
much later.
In view of the evidence compiled in the preceding paragraphs, a
feasible explanation of the evolution of the nymphaloid butterflies
seems to require serious consideration of tropical America as some-
thing other than a receptacle for pre-existing Holarctic groups. Prob-
ably a relatively undifferentiated nymphaloid stock either entered the
Neotropics as part of the extensive Holarctic invasion during the
MEM. AMER. ENT. SOC, 24
134 THE SATYRIDAE
Cretaceous (Darlington, 1957: 561-564), or this stock was already
in tropical America prior to the connection of South America with
the Holarctic. Since there are so many highly primitive groups of
the Satyridae, Morphidae, Ithomiidae, and Danaidae in the Neo-
tropics, it further appears that it was in tropical America that the
fundamental bifurcation of the nymphaloid stock occurred. At least
these four families appear to have arisen in the Neotropics. Later
all four reinvaded the Holarctic, probably during the Cretaceous, but
only two of them (Satyridae and Danaidae) reinvaded the American
tropics during the Tertiary. The Tertiary history of the danaids
probably parallels that of the satyrids, which will be discussed later.
Endemic, primitive, though often highly specialized, stocks of all
four families may be still found in the Neotropics. It seems most
likely, therefore, that these families all arose in the Cretaceous (pos-
sibly earlier) rainforests of tropical America.
The Later History of the Satyridae
The most primitive satyrids, the Haeterinae, are now restricted
to the Neotropical rainforest, and there is no indication that they
ever were elsewhere, and all evolution in this group has occurred
within the American tropical forests (Fig. 316). No members en-
tered the Old World, as did members of the Ithomiidae. It is also
doubtful that these insects ever exploited the grasslands that resulted
from progressive drying during the Tertiary. In fact, these insects
are singularly well adapted to the deep forest — the hyaline wings of
many species and the ghost-like fluttering flight render them virtually
invisible except when they pass through patches of sunlight. Masters
(in lift.) says that Haetera always restricts itself to the deepest forest,
never entering sunlit areas, but I have seen species of Pierella and
Callitaera sunning themselves early in the morning, but not later
in the day.
Figs. 317, 318. Fig. 317. The origin and subsequent distributional his-
tory of the Melanitini (Biinae) and the origin of the Elymniinae from the
Melanitini. For details, see text. Fig. 318. The origin and proliferation of
the Elymniinae into the tribes Lethini (see Fig. 319), Elymniini (Fig. 320),
Mycalesini (Fig. 321) and Zetherini. The latter tribe did not significantly
leave the Indo-Malayan region. The probable origin of the Eritinae is also
shown on this map. For details, see text.
LEE D. MILLER
135
ftethini (Fig. 319)
Elymniini(Fig. 320)
Elymniinae /
1 \\ Mycalesini (Fig. 321)
>•< V 7etherini
MEM. AMER. ENT. SOC. 24
136 THE SATYRIDAE
The history of the Brassolinae probably closely parallels that of
the Haeterinae (Fig. 316). These butterflies, however, are strong
fliers and have evolved a "flash coloration pattern" on the upper
surface and a cryptic under surface. They do not depend so much
on camouflage as on the strength of their flight. Many species are
crepuscular, particularly those occurring in more open country, but
most species are more or less restricted to the rainforest.
The Biinae did move into the Holarctic and apparently provided
the basis for the evolution of the rest of the family. Those biines
which did not leave the Neotropics (Fig. 316) have remained rain-
forest butterflies with patterns and flight characteristics similar to
those of the brassolines.
Whether the Neotropical members of any of the preceding sub-
families ever invaded the grasslands is questionable. If they did,
they were wiped out, probably in competition with the better adapted
Satyrinae which later came into the New World. It seems more
likely, however, that these butterflies never left the rainforest of
their origin and evolved into such ecologically successful species that
they easily resisted competition from the satyrines; perhaps these pri-
mitive groups actually forced the satyrines into more open country.
Those Biinae which did move into the Holarctic were obviously
much more like Antirrhini than Bia, but there are now no New World
antirrhines that have extremely close relationships with the Old World
species. The migrating biines were forced into the Paleotropics at a
fairly early time, where they apparently evolved into the present-
day tribe Melanitini ( Fig. 317). Several of the melanitines have
become very widespread, apparently relatively recently, and one,
Melanitis leda (Linne), is now found from West Africa to Mada-
gascar and east into Australia and Japan. Most of the species are
found in the Indo-Malayan region and in tropical Africa (Table 7).
The Melanitini responded to the increase in aridity of the early
Tertiary by invading open woodlands and even more open country,
being thus the first satyrids, historically and evolutionarily, to leave
the deep forest. These butterflies are more or less crepuscular, like
the Brassolinae. A possible explanation for the crepuscular flight
Figs. 319, 320. Fig. 319. The distributional history of the Lethini. See
text for details. Fig. 320. The distributional history of the Elymniini. See
text for details.
LEE D. MILLER
137
MEM. AMER. ENT. SOC, 24
138
THE SATYRIDAE
Fig.
details.
321. The distributional history of the Mycalesini. See text for
habit is that such species are actually approximating the conditions
of the deep forest as regards light intensity. In short, such species
are basically forest species that are "carrying the forest environment
with them". The successful invasion of the open country provided
the impetus for the great adaptive radiation and phylogenetic diversi-
fication of the satyrids in the Old World. The Neotropical Antirrhini,
on the other hand, never became successfully adapted to open country,
and without leaving the forests evolved into the present tribe with
fewer than twenty species. In the Indo-Australian region the Melani-
tini were probably giving rise, meanwhile, to the Elymniinae through
steps resembling the present-day genera Parantirrhoea (Melanitini),
Pty chandra and Samanta (both Lethini), shown in Figs. 317, 318.
The Elymniinae proliferated rapidly during late Cretaceous and
early Tertiary into the four tribes, most of which stayed in the wood-
lands and scrublands: some (a number of Mycalesini) invaded the
savannas, and others (the Neorina-series) re-entered the deep forests.
The pattern of early and later migration of the subfamily rather closely
parallels that of many vertebrate groups and seems to have been
LEE D. MILLER 139
limited northward by the Tethys Sea. Little movement from the
Paleotropics seems to have been accomplished before the closing of
this "water gap" between the Paleotropics and the Palearctic (Fig.
318). Interrelationships within the Elymniinae can be better drawn
than those within any other subfamily. The Lethini grade toward the
Zetherini, on the one hand, and the Mycalesini (through such genera
as Mandarinia and Orsotriaena), on the other, as brought out in
the systematic revision. The Mycalesini connect well with the Elym-
niini, as do the zetherines. The elymniines are "throwbacks'"' in one
respect: all known members feed as larvae on Palmaceae, hence the
common name "palmflies". All tribes of the Elymniinae but the
Zetherini are widely distributed in the Old World at present, although
only one — the Pararge-section of the Lethini — has managed signifi-
cantly to penetrate the Palearctic. The distributional histories of the
tribes are shown in Figs. 319 (Lethini), 320 (Elymniini), 321
(Mycalesini); the actual patterns are discussed in the analyses of the
individual faunae of the world.
Apparently within a short time — probably still in the Cretaceous
— the Elymniinae gave rise to the Satyrinae (Figs. 318, 322), pre-
sumably through either the Mycalesini or (more probably) the
Lethini and the Hypocystini of the Satyrinae. It is also probable that
the Eritinae arose from a basic elymniine stock (Fig. 318) and pos-
sible that the Ragadiinae did (Fig. 322), but the origin of the latter
subfamily is among the most obscure in the family Satyridae.
The hypocystine genus Lamprolenis, while a fairly typical member
of that tribe, retains some very primitive characteristics similar to
those of the Mycalesini (Fig. 148). There is a typical Mycalesis-
like hair tuft on the hindwing, and the general facies embody a com-
bination of mycalesine and hypocystine characteristics.
The other case of similarity between the Elymniinae and Satyrinae
is to be found in South Africa, where members of the Aeropetes-series
of the Lethini seem to grade into the Dirini (Figs. 67-72 and 248-259,
respectively). It would be tempting to ascribe great evolutionary
significance to the similarity, but the A eropetes-series contains primi-
tive lethines, whereas the Dirini appear to be fairly advanced Sa-
tyrinae — particularly as regards the reduction of the forelegs of both
sexes. Therefore, this similarity is ascribed to convergence, not to
phyletic proximity.
MEM. AMER. ENT. SOC, 24
140 THE SATYRIDAE
The strongest link between the Elymniinae and the satyrines
occurs through the subfamily Eritinae. Eritines have many charac-
teristics in common with the Lethini, as discussed in the systematic
revision, and also share characters with the Hypocystini. This can-
not be merely convergence, and it appears that the Satyrinae arose
in the Indo-Australian region from an elymniine stock not far re-
moved from the lethine stock (Figs. 318, 322).
In any event, the Satyrinae — probably from a Hypocystini-like
stock — soon split into two basic groups, the Ypthimini- and Satyrini-
sections (Fig. 322). This dichotomy was probably brought about
by the increasing aridity and proliferation of savannas during the
early Tertiary. The Ypthimini-section generally occupies more open
country than do the Satyrini-section butterflies.
During the early to mid-Tertiary the various tribes of the Satyri-
nae, except the Pronophilini and Euptychiini, probably evolved in
the Old World (Figs. 323-326). Upon the reopening of the Cen-
tral American land connection between North and South America —
it was probably sufficiently opened in Miocene, perhaps Oligocene,
times — the forerunners of the principal South American tribes of
the Satyrinae, the Pronophilini (derived from the Satyrini-section,
Fig. 325) and the Euptychiini (from the Ypthimini-section, Fig.
323 ) reentered South America and rapidly assumed dominance over
the "old" South American fauna. At least part of this dominance
may be ascribed to the ecological diversity shown by both tribes, as
explained in the discussions of the tribes in the systematic revision.
A glance at the numbers of species involved in both the Pronophilini
and Euptychiini shows the dramatic radiation that has taken place in
the Neotropics.
It is doubtful that the radiation of the Euptychiini and Pronophi-
lini resulted in a wholesale extinction of the Haeterinae, Brassolinae
and Biinae, since the new arrivals probably did not compete ecologi-
cally with the already established primitive ones. Both the Pro-
Figs. 322, 323. Fig. 322. The origin of the Satyrinae from the Elymniinae
and the division of the satyrines into the Hypocystini, the Ypthimini-section and
the Satyrini-section. The probable origin of the Ragadiinae from the Elym-
niinae is also shown in this figure. For details, see text. Fig. 323. The origin
and distributional history of the Ypthimini (Old World) and Euptychiini (New
World). See text for details.
LEE D. MILLER
141
Ypthimin i-sect_
Satyrini-sect.f Satyrinae
o
elymniine ancestor
Ragadiinae
Hypocystini
MEM. AMER. ENT. SOC, 24
142
THE SATYRIDAE
Fig. 324. The origin and distributional history of the Coenonymphini,
Maniolini and Erebiini. See text for details.
nophilini and Euptychiini feed as larvae predominantly on Grami-
neae, which are probably not used extensively by the primitive species.
Just before and during the Pleistocene there was an exchange
of faunal elements between the Nearctic and the Palearctic, resulting
in such Holarctic distributions as those of Coenonympha (Fig. 324),
Oeneis (Fig. 326) and Erebia (Fig. 324), although these genera
probably differentiated much earlier, and some of the Holarctic mem-
bers of the Maniolini (Fig. 324) and Satyrini (Fig. 326).
A proposed phylogeny for the Satyridae is presented in Fig. 327,
embodying the information given above.
Derivation of the Satyrid Faunae of the World
Comparisons of the faunal compositions of various parts of the
world often prove valuable in determining the derivation of the
elements of some areas. With this assumption in mind Table 7 was
drawn to show the composition by tribes of the Satyridae in major
faunal regions of the world.
LEE D. MILLER
143
Figs. 325, 326. Fig. 325. The origin and distributional history of the
Dirini and Pronophilini. See text for details. Fig. 326. The origin and dis-
tributional history of the Satyrini and Melanargiini. See text for details.
MEM. AMER. ENT. SOC, 24
144
THE SATYR1DAE
Cretaceous
TIME approx.
early Tertiary
TAXON
mid-Tertiary
Tribe
Subfamily
Melanargiini _
Satyrini
Ptonophilini
Ditlni
Hypocystini
Erebiini
Maniolinl
Coenonymphini
Ypthimini
Euptychiini ''
>SATYRINAE
Eritlni
Mycalesin J-___
Ely m niini
Zetherini
Lethini- —
Melanitini
Antirrhini
Biini
Bfassolini
Haeterini
-RAGADIINAE
— ERITINAE
_~~~-ELYMNIINAE
_.;::: BIINAE
-BRASSOLINAE
--HAETERINAE
Fig. 327. A proposed phylogeny of the Satyridae showing the relation-
ships of the tribes. The time scales are very approximate and conform to the
hypotheses proposed in the section on evolution and zoogeography. For details,
see text.
LEE D. MILLER 145
The Neotropics. — The Neotropics, as broadly and somewhat incor-
rectly defined to include Antarctic, temperate and tropical regions
of South America, show a satyrid fauna composed of six tribes which
are members of four subfamilies. Five of these tribes are endemic,
and four of them — Haeterini, Brassolini, Antirrhini and Biini — are
the most primitive in the family. The fifth endemic tribe, the Pro-
nophilini, is a member of the Satyrinae and most closely related to
the Holarctic Satyrini. The pronophilines probably reached, during
the mid-Tertiary, South America and proliferated there. The Eup-
tychiini are exclusively American satyrids most closely allied to the
Ypthimini. They were probably derived from a common stock with
the Ypthimini, introduced into the Neotropics and proliferated there
contemporaneous with the Pronophilini. The euptychiines, while
entirely American and overwhelmingly Neotropical, cannot be con-
sidered an endemic tribe because seven species are found in the
Nearctic, as far north as southern Canada.
In summary the Neotropical satyrid fauna appears to have been
derived from two sources. The Haeterinae, Brassolinae, Antirrhini
and Biini evolved in tropical America from the basic satyrid stock
which either arose in tropical America from a basic nymphaloid stock
that was already there or evolved from such a stock that invaded
tropical America during the Cretaceous. The Neotropics were then
reinvaded, probably in the Oligocene, perhaps even in the Eocene,
by two basically Paleotropical groups, the Pronophilini (derived
from a Satyrini-section stock) and the Euptychiini (from a Ypthimini-
section stock). There has been no extensive immigration in recent
times from the Nearctic; indeed, most recent movement in the Amer-
icas of the Satyridae has been northward from the Neotropics.
The Nearctic. — The Nearctic fauna has been derived primarily
from the Neotropics and the Palearctic. Seven tribes are represented
in North America, none endemic, and most species are very recent
arrivals, many during the Pleistocene. By far the majority of the
species in the Nearctic are assignable to Holarctic tribes or genera.
A few of these, such as Neominois ridinsii, Enodia portlandia and
creola, Satyrodes eurydice, Coenonympha haydenii and Cercyonis
species, appear to be pre-Pleistocene arrivals; these butterflies prob-
ably arrived in North America in the Miocene or Pliocene, perhaps
even earlier. The remainder of the genera arriving from the Palearctic
— Oeneis, most Coenonympha, Erebia — belong to a basically arctic-
MEM. AMER. ENT. SOC, 24
146 THE SATYRIDAE
Table 7
The taxonomic composition of the world satyrid fauna. a= single species;
b = 2 to 5 species; c = 5 to 10 species; d = more than 10 species).
Taxon
o
'a,
o
u>
o
u
z
u
o
o
ca
a
Oh
W
c
—
6
C
c
•—
<
o
<
o
H
OS
_o
<
d
6
m
H
a
o
Hi
a
60
a
Haeterinae
d
Brassolinae
d
Biinae
d
a
d
a
c
b
b
Antirrhini
d
Biini
a
Melanitini
a
d
a
c
b
b
Manataria
b
Elymniinae
b
d
C
d
d
d
d
d
Lethini
b
d
c
d
a
b
Zetherini
a
b
Elymniini
d
b
b
b
a
Mycalesini
c
d
d
d
c
d
Eritinae
b
Ragadiinae
b
c
Satyrinae
d
d
d
d
d
d
d
d
a
Hypocystini
d
Ypthimini
d
b
d
b
d
d
a
Palaeonympha
a
Euptychiini
d
c
Coenonymphini
b
d
d
Maniolini
b
d
d
Erebiini
c
d
d
Dirini
c
Pronophilini
d
a
Satyrini
d
d
d
b
Melanargiini
d
d
Endemic tribes
5
1
2
1
1
alpine fauna and probably date from the Pleistocene: Coenonympha
mixturata found in the bogs of Alaska and the Yukon is virtually
inseparable from the Siberian population and must be a post-Pleisto-
cene immigrant. The tribes derived from the Palearctic are the
Lethini, Coenonymphini, Maniolini, Erebiini and Satyrini. Two
LEE D. MILLER 147
tribes, the Pronophilini and Euptychiini, have entered the Nearctic
from the Neotropics. The first of these tribes is represented by a
single species, Gyrocheilus patrobas, which reaches southern Arizona.
This invasion is probably of rather recent origin, perhaps post-
Pleistocene, though it is difficult to assign dates to an invasion of a
single species. The Euptychiini have left several species in North
America, and one group — the Megisto cymela complex — has become
rather distinct from its Neotropical progenitors. The cymela-gvoup
probably arrived well before the Pleistocene, whereas the other species
in the Euptychiini are closely allied to tropical members and may
have arrived during an interglacial.
The Palearctic. — The importance of the Palearctic, while it was
not the place of the origin of the Satyridae, nevertheless cannot be
overstressed with respect to subsequent evolution. Eleven tribes,
representing four subfamilies, are found in this region, although the
"true" Palearctic fauna is composed only of the Elymniinae and the
Satyrinae. The other two subfamilies, the Biinae and the Ragadiinae,
enter the Palearctic only in its southeastern part, were derived from
the Indo-Malayan region and need be mentioned only in passing.
The Elymniinae are represented by the tribes Lethini, Zetherini
and Mycalesini — the last two being found just in the southeastern
part of the area and also properly are part of the Indo-Malayan fauna.
The tribe Lethini is represented chiefly by the Pararge-senes — more
boreal in its distribution than other lethine groups — which probably
evolved in the mid-Tertiary and became cold-adapted. The inter-
esting link between the Lethini and Mycalesini, Mandarinia regalis,
is found in temperate China, suggesting that the latter tribe may once
have been found much farther north than it is today.
The Palearctic is the metropolis of most of the tribes of the
Satyrinae, with six represented. Five of these tribes — Satyrini, Me-
lanargiini, Coenonymphini, Maniolini and Erebiini — probably arose
in the Palearctic itself in response to progressive drying, with the
increase of grasslands, during the early Tertiary. Only the Melanar-
giini are truly endemic, all other tribes having expanded into the
Nearctic during the later Tertiary or Quaternary. The sixth satyrine
tribe, the Ypthimini, is primarily Paleotropical, and the Palearctic
ypthimines must have been derived from the tribes. One group within
the ypthimines, the Callerebia-series, is exclusively Palearctic and
probably arose from the ypthimine stem about the same time as the
MEM. AMER. ENT. SOC, 24
148 THE SATYRIDAE
other tribes mentioned above. All members of the Callerebia-sevies
now are found under the stringent ecological conditions of the central
Asian mountains; none has invaded the main part of the region.
In sharp contrast with the Nearctic, the Palearctic has been very
important in the later evolution of the Satyridae. Just why the Pale-
arctic should have played such a role in the evolution of the Satyrinae
is a puzzle: there were certainly vast savannas in the Nearctic during
the Tertiary (Darlington, 1957), which proved to be the route of
dispersal of the forerunners of the Pronophilini and Euptychiini and
thus must have been occupied during the Tertiary. The primitive
forms of the five Holarctic satyrine tribes are all Palearctic, not
Nearctic. The evidence for these tribes having arisen in the Old
World is overwhelming. The only tribe which is a candidate for
arising in part in the Nearctic is the Satyrini: Neominois ridingsii is
a rather primitive species, most nearly related to the Asian Karanasa.
The great proliferation of the Satyrus-series (Satyrini) in central
Asia argues strongly for its being considered the "birthplace" of the
tribe. In all fairness, however, it must be pointed out that North
America was the place of the greatest Tertiary development of the
horses and camels (Darlington, 1957: 352-354), yet no native
species of either group is presently found in North America. Such
an evolutionary pattern may explain the satyrids — evolution of some
stocks in the New World, spread of these stocks into the Old World,
extinction of the Nearctic fauna and reinvasion and repopulation of
the Nearctic by Palearctic groups. In the absence of a fossil record, I
must accept the neontological evidence that the satyrine tribes evolved
in the areas of their present greatest density and diversity. Parallels
exist: Simpson (1953: 51-52) mentions the New World oreodonts
and pronghorn "antelopes" and the Old World giraffes. None of
these groups left the hemisphere of their origin.
The Indo-Malayan region. — This area, bounded roughly by India,
the Philippines and Wallace's line, has been likened to the "Garden
of Eden" by generations of zoogeographers. Its satyrid fauna is
strong only in moderately primitive groups. In all, nine tribes of
five subfamilies are represented in the Indo-Malayan fauna. This is
the area of the best development of the Ragadiinae and Eritinae, and
these are to be considered truly endemic to the region. The melani-
tines (Biinae) are best represented in the Indo-Malayan fauna and
probably arose there or arrived at an early date.
LEE D. MILLER 149
The Elymniinae have reached their highest development in this
region, where the four tribes are best represented; all probably arose
there. Those genera tending to connect tribes, with the exception of
Mandarinia, are found on the Malay Peninsula. The bifurcation of
the Mycalesini into naked and hairy eyed species took place in this
region, and both groups are still found throughout the Indo-Malayan
region. These have spread and provide some rather interesting data
in the analysis of the African-Malgache fauna. Certain considera-
tions, outlined in the systematic revision, led to the conclusion that
the Lethini are the most primitive elymniines. Most lethines have
hairy eyes, and for this reason the hairy-eyed condition is considered
primitive in the Mycalesini (see also the discussion under the Mal-
gache fauna, which follows).
The Satyrinae are represented by two tribes, the Satyrini and
Ypthimini. The former is a secondary immigrant from the Palearctic,
only entering the Indo-Malayan region along its northern end. The
Ypthimini, however, probably arose in this region and apparently
replaced its immediate relatives — forms closer to the present-day
Hypocysta-series (Hypocystini), which is now restricted to the Aus-
tralian region east of Wallace's line. The ypthimines spread during
the early Tertiary into Africa and the New World, giving rise to the
Euptychiini of the latter region.
In summary, due to the prevalence of the Melanitini, Elymniinae
and intermediate genera in this region, the Indo-Malayan region is
considered to have given rise to the elymniines. The Eritinae and
Ragadiinae also arose in this area. The Indo-Malayan region did
not play an important role in the later evolution of the Satyrinae: the
focus of that subfamily shifted to the Palearctic.
The Australian Region (including the Pacific Islands). — Faunistically
the Australian region, including Australia, New Guinea, New Zea-
land and the nearby islands, is one of the most interesting. It is
distantly related to the Indo-Malayan region, but substantial inter-
change of faunal elements between these regions occurred only in the
distant past. Simpson (1961b: 443) has dated the extensive immi-
gration of the marsupials into Australia as Cretaceous or Paleocene,
and it was probably at that time that much of the regional butterfly
fauna arrived. Elsewhere Simpson (1961b: 435-436) shows that
several waves of rodent groups arrived in the Australian region from
MEM. AMER. ENT. SOC, 24
150 THE SATYRIDAE
the Indo-Malayan from Miocene time to the present. All the tribes
represented in the Australian region have also penetrated the Pacific
islands, over water, hence the islands are considered along with the
continent.
Just one tribe is endemic in this region, the Hypocystini. The
metropolis of the Xenica-series is Australia, with some aberrant spe-
cies in New Zealand. All the satyrids in New Zealand were derived
long ago from the Australian ones. The Hypocysta-series is best
developed in New Guinea and has spread into the Pacific islands.
The other satyrine tribe represented in this region is the Ypthimini,
the members of which are highly modified, particularly the Pacific
island genus Xois. The Ypthimini, as shown by their expansion into
Africa and the Americas, was a fairly plastic group.
A single Melanitis is found throughout the region in question,
M. leda, a widespread species distributed west to the westernmost
part of Africa. The same may be said about those Elymniini which
occur on these islands; all are widespread species. The arrival of
Melanitis and Elymnias must have been a recent event since they are
little differentiated from their continental progenitors. The Myca-
lesini of the region reflect the probably early spread of this tribe, since
it also reached Madagascar.
In summary, the satyrid fauna of the Australian region was
derived from the Indo-Malayan region, and this derivation very
early, resulting in a highly distinctive fauna. If the time schedule
for the marsupials is valid for the satyrids, most of the groups reached
the region in late Cretaceous to earliest Tertiary times. This indi-
cates that the Hypocystini of the most evolved subfamily, Satyrinae,
were in existence by the earliest Tertiary. A few strong flying species,
such as M. leda, probably arrived over water very recently.
The Ethiopian Region. — This region includes both the tropical part
of Africa and temperate South Africa. These are faunistically very
different areas, but they are so intimately related that they must be
considered together.
There is a single endemic African satyrid tribe, the Dirini. This
tribe, which has morphological affinities with both the Elymniinae
and the Satyrini-section of the Satyrinae, is restricted to South Africa.
Several interpretations are possible for the Dirini: (1) the Satyrinae
arose in Africa, (2) the Dirini were exterminated in all places but
LEE D. MILLER 151
South Africa or (3) the Dirini stock was isolated in South Africa
with the expansion of the tropical rainforest belt into eastern Africa,
and it evolved there. The first alternative is highly improbable, as
I have shown in the discussion of the Hypocystini in the systematic
revision, but either (2) or (3) might be possible. In the absence of
a fossil record we shall never really know. The only other tribe of
the Satyrinae represented in the African fauna is the Ypthimini. The
ypthimines of tropical Africa are closely related to the Indo-Malayan
ones, whereas most of the South African ypthimines belong to the
more or less endemic Melampias-series, one member of which is also
found on Madagascar. It seems likely that the Melampias-series
were early invaders of Africa — contemporaneous with the invasion
of the Dirini — which differentiated and were then forced south and
were replaced in tropical Africa by the ypthimines that arrived later.
The Lethini show a similar pattern. In southern Africa two
very primitive lethines, the A eropet es-series, are found which prob-
ably are remnants of an early fauna. The East and South African
genus Aphysoneura is much more like the Indo-Malayan lethines,
and it was probably a later faunal addition. The Mycalesini have
made a significant invasion of Africa. All the species on the conti-
nent have naked eyes, unlike those on Madagascar, and probably
represent a later immigration, contemporaneous with the arrival of
the "conventional" ypthimines and Aphysoneura. The mycalesines
will be considered in further detail in the discussion of the fauna of
Madagascar. The elymniines and the melanitines of Africa are rep-
resentatives of the most widespread groups and almost certainly
arrived from the Indo-Malayan region, probably fairly recently:
frequently the African members are conspecific with their Indo-
Malayan counterparts.
In summary, Africa appears to have been populated more than
once from the Indo-Malayan region. The tropical African satyrid
fauna looks more like a depauperate Indo-Malayan fauna than any-
thing else, but that of temperate Africa has evolved significantly and
is quite distinct. Probably the South African endemic groups rep-
resent the "old" African fauna of the early Tertiary, whereas that
of tropical Africa is younger, perhaps dating from the Oligocene or
even the Miocene (Darlington, 1957: 590). Such species as Mela-
nitis leda probably arrived in tropical Africa much more recently.
MEM. AMER. ENT. SOC, 24
152 THE SATYRIDAE
Madagascar. — There can be little doubt that the colonization of
Madagascar was from Africa, but mostly so long ago that the species
bear little resemblance to species now found on continental Africa.
The Melanitini and the Elymniini of this island, on the other hand,
are conspecific with their mainland counterparts, as is the only mem-
ber of the Satyrinae, a species of Ypthimini: all these butterflies are
probably recent introductions, no doubt by waif dispersal.
The most interesting satyrids in Madagascar, from a zoogeo-
graphic standpoint, are the Mycalesini — the dominant tribe on the
island. As noted before, both naked- and hairy-eyed mycalesines
occur in the Indo-Malayan regions, whereas those from continental
Africa have naked eyes. The species found in Madagascar, without
exception, have hairy eyes. One would be tempted at first glance to
postulate that the Malgache mycalesines were derived directly from
the Indo-Malayan ones. This is precisely why the mythical continent
of "Lemuria" was postulated: a land bridge to account for the similar
Malgache and Indo-Malayan lemurs; the Madagascar genera are less
like continental African genera than any of the continental genera
are like each other. In the mycalesines, dissimilarities indicate a long
isolation of the Malgache fauna. For this reason it seems more
likely that the mycalesines migrated into Africa very early (Creta-
ceous or Paleocene?), and that stock which made the first immigra-
tion was hairy-eyed, as is characteristic of primitive elymniines. Fur-
thermore, the Malgache mycalesines have a less clubbed female
foretarsus than do the continental species. These mycalesines were
contemporaneous with the Dirini, Melampias-series of the Ypthimini
and the Aeropetes-series of the Lethini and formed part of the "old"
African satyrid fauna. Madagascar received its mycalesine fauna
from this very early invasion. Just when this occurred is problematic:
Matthew (1915: 204-205) thought that Madagascar and Africa
were never physically connected, and Darlington (1957: 535) be-
lieved that the Malgache fauna may have begun to accumulate in the
Oligocene. In any event, the Malgache mycalesine fauna is probably
of early Tertiary age. Climatic changes or pressure of competition by
the later mycalesine immigrants — which did not reach Madagascar —
replaced the hairy-eyed mycalesines on mainland Africa, leaving the
morphologically primitive Malgache fauna completely isolated geo-
LEE D. MILLER 153
graphically from their nearest relatives in the Indo-Malayan region.
Such invasion and subsequent extermination of geographically inter-
mediate populations is well known among fossil vertebrates (Dar-
lington, 1957: 102-121; 159-172; 206-230).
Bibliography
Agenjo, R. 1947. Catalogo ordenador de los lepidopteros de Espana. Gra-
ellsia, 5(3): unnumbered pages at end.
d'Almeida, R. F. 1951. Ligeiras observacoes sobre o genero Cithaerias
Hubner. 1819 (Lep Satyridae). Arq. Zool. Sao Paulo. 493-505; 2 pis.
Aurivillius, C. 1882. Recensio critica Lepidopterorum Musei Ludovicae
Ulricae, quae descripsit Carolus A. Linne. K. Svenska Akad. Handl.
19(5): 3-188: ill.
Bang-Haas, O. 1938. Neubeschreibungen und Berichtigungen der palaeark-
tischen Macrolepidopterenfauna. Ent. Zeitg. Frankfurt 52: 177-180.
Billberg, G. J. 1820. Enumeratio Insectorum in Museo G. J. Billberg.
Stockholm, Priv. publ.: ii + 138 pp.
Blanchard, C. E. 1840. Lepidopteres. in Castelnau, F. L., Histoire Naturelle
des Insectes. . . Avec Introduction . . . par M. Brulle. Paris, vol.
3: 417-560.
. 1852. Lepidoptera. ;'/; Gay, C., Historia fisica y politica de
Chile. . . Paris and Santiago, Priv. publ.: vol. 7.
Boisduval, J. B. A. D. 1832. Faune entomologique . . . Pt. 1, Lepidopteres.
in Voyage . . . de FAstrolabe . . . pendant 1826-1829. Paris, J. Tatsu:
i-iv: 5-267 pp.; ill.
. [1 833]( 1832-1843). Icones historique des Lepidopteres d'Eur-
ope. . . 1. Rhopaloceres. Paris, Priv. publ.: 1-251; pis. 1-47.
. 1836. Histoire naturelle des Insectes. Species general de Lepi-
dopteres. Paris, Libr. Encycl. de Roret: xii + 690 pp.; ill.
. 1870. Considerations sur des Lepidopteres envoyes du Guate-
mala a M. de l'Orza. Rennes, France, Oberthiir et Fils: 100 pp.
Brown, F. M. 1941. Some notes on four primary reference works for Lepi-
doptera. Ann. Ent. Soc. America 34(1): 127-138.
. 1942. Notes on Ecuadorian butterflies, V. Jour. New York
Ent. Soc. 50: 309-333; ill.
. 1944. Notes on Ecuadorian butterflies, IV. The genus Penro-
sada, new. Ann. Ent. Soc. America 37(2): 255-260.
Brown, W. L., Jr. 1965. Numerical taxonomy, convergence and evolutionary
reduction. Syst. Zool. 14(2): 101-109; ill.
Butler, A. G. 1867a. Remarks on the nomenclature of the European genera
of satyridae Lepidoptera. The Entomologist 3: 277-281.
mem. amer. ent. soc, 24
154 THE SATYRIDAE
1867b. Description of a new genus and species of diurnal
Lepidoptera. Ent. Mo. Mag. 4: 121-122; ill.
. 1867c. Description of a new genus of diurnal Lepidoptera
belonging to the family Satyridae. Ann. Mag. Nat. Hist. (3)19:
49-51; ill.
. 1867d. Descriptions of five new genera and some new species
of satyride Lepidoptera. Ann. Mag. Nat. Hist. (3)19; 161-167; ill.
. 1867e. Descriptions of some remarkable new species and a
new genus of diurnal Lepidoptera. Ann. Mag. Nat. Hist. (3)20:
216-217; ill.
1867f. Revision of the group of lepidopterous insects hitherto
included in the genus Pronophila of Westwood. Ann. Mag. Nat. Hist.
(3)20: 266-268; ill.
1867g. Description of a new genus and species of American
Satyridae from the collection of Mr. H. W. Bates. Ann. Mag. Nat.
Hist. (3)20: 404-405; ill.
. 1868a. An essay towards an arrangement of the genera of the
family Satyridae. Ent. Mo. Mag. 4: 193-197.
. 1868b. Catalogue of the diurnal Lepidoptera of the family
Satyridae in the collection of the British Museum. London, Trustees
of the British Mus.: vi + 211 pp.; ill.
1871. Descriptions of five new species, and a new genus, of
diurnal Lepidoptera, from Shanghai. Trans. Ent. Soc. London 19:
401-403.
. 1872. Descriptions of new butterflies from Costa Rica. Cis-
tula Ent. 1 : 72-90.
. 1874. Description of a new genus and species of satyridian
butterflies. Ent. Mo. Mag. 10: 204-205.
. 1876. List of the butterflies now known to inhabit New Zea-
land, with descriptions of a new genus, and a new species, in the col-
lection of John D. Enys, Esq. Ent. Mo. Mag. 13: 152-154.
1879. Descriptions of new species of Lepidoptera from Mada-
gascar, with notes on some of the forms already described. Ann. Mag.
Nat. Hist. (5)4: 227-246.
. 1880. On a collection of Lepidoptera from Madagascar, with
descriptions of new genera and species. Ann. Mag. Nat. Hist. (5)5:
333-344.
. 1884. On a new genus of butterfly from New Zealand. Ann.
Mag. Nat. Hist. (5)13: 171-173: ill.
1885. An account of two collections of Lepidoptera recently
received from Somali-land. Proc. Zool. Soc. London [1885]: 756-776;
ill.
. 1899. On a collection of butterflies almost entirely made at
Salisbury, Mashunaland, by Mr. Guy A. K. Marshall in 1898. Proc.
Zool. Soc. London [1898]: 902-912.
LEE D. MILLER 155
Carpenter, F. M. 1930. A review of our present knowledge of the geo-
logical history of insects. Psyche 37: 15-34.
Caster, K. E. 1952. Stratigraphic and paleontologic data relevant to the
problem of Afro-American ligation during the Paleozoic and Mesozoic.
in Mayr, E., 1952: 105-152.
Clark, A. H. 1947. The interrelationships of the several groups within the
butterfly superfamily Nymphaloidea. Proc. Ent. Soc. Washington
49(6): 148-149.
. 1948. Classification of the butterflies, with the allocation of
the genera occurring in North America north of Mexico. Proc. Biol.
Soc. London 61: 77-84.
Condamin, M. 1960. Distinction de deux sous-genres dans le genre Myca-
lesis (Lep. Satyridae). Bull. Inst, franc. d'Afrique Noire (A) 22(4):
1252-1258.
. 1961. Mises au point de synonymie et descriptions de nouveaux
Bicyclus (Lep. Satyridae). Bull. Inst, franc,. d'Afrique Noire, (A) 23-
(3): 782-799; ill.
Dalman, J. W. 1816. Forsok till systematisk Uppstallning af Sveriges Fja-
rilar. K. Svenska Akad. Handl. 37: 48-101, 199-225.
Darlington, P. J. Jr. 1957. Zoogeography: the geographical distribution of
animals. New York, John Wiley and Sons: vii-xi + 675 pp.; ill.
Doherty, W. 1886. A list of butterflies taken in Kumaon. Jour. Asiatic
Soc. Bengal 55: 103-140.
Doubleday, E. 1845. Descriptions of new or imperfectly described diurnal
Lepidoptera. Ann. Mag. Nat. Hist. (1)16: 304-308.
. 1848a. in Hewitson, W. C, Description of a new genus and
species of Satyridae. Proc. Zool. Soc. London [1848]: 115-117.
. 1848b. List of the specimens of lepidopterous insects in . . .
the British Museum. London, Trustees of the British Mus.: 37 pp.
Doubleday, E., J. O. Westwood and W. C. Hewitson 1846-1852. The
genera of diurnal Lepidoptera. . . 2 vols. London, Longman, Brown,
Green and Longman's: 534 pp.; ill.
Doumet, N. 1861. Description de Lepidopteres nouveaux. Revue Zool.
(2)13: 171-178; ill.
Dunbar, C. O. 1952. Discussion, in Mayr, E., 1952: 153-158.
Duncan, J. 1837. Foreign butterflies. London, The Naturalist's Library
(No. 18): 180 pp.
[Dunning, J. W. and Pickard] 1858. An accentuated list of the British Lepi-
doptera with hints on the derivation of the names; published by the
Entomological Society of Oxford. London, Van Voorst: 44 + 118 pp.
Ehrlich, P. R. 1958. The comparative morphology, phylogeny and higher
classification of the butterflies (Lepidoptera: Papilionoidea). Univ.
Kansas Sci. Bull. 39(8): 305-370: ill.
MEM. AMER. ENT. SOC, 24
156 THE SATYRIDAE
Ehrlich, P. R. and P. H. Raven. 1965. Butterflies and plants: a study in
coevolution. Evolution 18(4): 586-608.
El wes, H. J. 1899. On the Lepidoptera of the Altai Mountains. Trans.
Ent. Soc. London 47: 295-367: ill.
Fabricius, J. C. 1807. (Title not available). Mag. fur Insecktenkunde
(Illiger) 6: 282-284 (fide Hemming, F., 1934).
Felder, C. 1861. Ein neues Lepidopteren aus der Familie Nymphaliden und
seine Stellung in natiirlichen Systeme, begriindet aus der Synopse der
iibrigen Gattungen. Nov. Act. Acad. Caesar Leopold, 28(3): 1-50.
Felder, C. and R. Felder. 1859. Lepidopterologische Fragmente. III. Wie-
ner Ent. Monatschr. 3:321-346.
. 1860. Lepidoptera nova in paeninsula Malayica collecta. Wie-
ner Ent. Monatschr. 4: 395-402.
1861. Lepidoptera nova a Dre. Carolo Semper in insulis Philip-
pinis collecta. Wiener Ent. Monatschr. 5: 297-306.
. 1867 (1864-1867). Reise der Osterreichischen Fregatte No-
vara. . . Zool. 2. Lepidoptera. Wien, Carl Gerold's Sohn: i-vi + 548
pp.; ill.
Felder, R. 1869. Diagnosen neuer von dem k. k. Oberlieutnant H. V. Hede-
mann in Mexico in Jahren 1865-1867 gesammelten Lepidopteren. Ver-
hdl. zool.-Bot. gesell. Wien 19: 465-480.
Fereday, R. W. 1879. Description of a new genus and species of butterfly
of the subfamily Satyrinae. Ent. Mo. Mag. 16: 128-130.
Forbes, W. T. M. 1932. How old are the Lepidoptera? American Nat.
66: 452-460.
Forster, W. 1964. Beitrage zur Kenntnis der Insecktenfauna Boliviens XIX.
Lepidoptera III, Satyridae. Veroff. Zool. Staatssamml. Miinchen 8:
51-188; ill.
Fox, R. M. 1956. A monograph of the Ithomiidae (Lepidoptera), part 1.
Bull. American Mus. Nat. Hist., Ill: 1-76; ill.
. 1967. Forelegs of butterflies I. Introduction: chemoreception.
Jour. Res. Lepid. 5(1): 1-12: ill.
Fox, R. M. and J. W. Fox 1964. Introduction to comparative entomology.
New York, Reinhold Publ. Co.: 450 pp.; ill.
Fruhstorfer, H. 1898. Neue Mycalesis aus Celebes. Stettiner Ent. Zeitg.
59: 263-266.
. 1907. Monographie der Elymniinae. Iris, 20: 157-252; ill.
. 1908. Neue indo-australische Mycalesis und Besprechung ver-
wandter Formen. Verhdl. Zool.-Bot. gesell. Wien 58: 126-239.
■ . 1909. Neue Satyriden. Internatl. Ent. Zeits., Giiben 3: 130-
131, 133-135.
1912 (1912-1915). Nymphalidae, etc. in Seitz, A., Der
Grossschmetterlinge der Erde, 9 (Indo-Australische Tagfalter). Stutt-
gart, Alfred Kernan Verlag: pp. 119-823:
LEE D. MILLER 157
Gaede, M. 1931. Satyridae. in Strand, E., (ed.), Lepidopterorum Cata-
logus, 29 (43, 46, 48). Berlin, W. Junk: 759 pp.
G erst acker, A. 1871. Beitrag zur Insecktenfauna von Zanzibar. Arch,
fur Naturgesch. (A)37(l): 345-363.
Godman, F. D. and O. Salvin. 1880. Descriptions of some supposed new
species of butterflies from New Guinea. Proc. Zool. Soc. London
[1880]: 609-614: ill.
Gray, G. R. 1846. Descriptions and figures of some new lepidopterous in-
sects, chiefly from Nepaul. London: 15 pp.
Grote, A. R. 1873. On Mr. Scudder's systematic revision of some of the
American butterflies. Paper No. 2. Canadian Ent. 5: 62-63.
Hampson, G. F. 1891. Lepidoptera from the Subaki River, East Africa,
with descriptions of new species. Ann. Mag. Nat. Hist., (6)7: 179-184.
. 1918. List of the families and subfamilies of the Lepidoptera.
Novitates Zool., 25: 383-394.
Hayward, K. J. 1949. Satiridos Argentinos nuevos para la Ciencia (Lep.
Satyridae). Acta Zool. Lilloana 8: 151-159.
. 1953. Satiridos Argentinos (Lep. Rhop. Satyridae). I. Los
generos (excluidos Euptychia y Neomaniola). Acta Zool. Lilloana
15: 5-66: ill.
Heimlich, W. 1959. Eine neue Satyride aus Chile. Ent. Zeits. (Stutt-
gart) 69: 173-179: ill.
Hemming, F. 1933. On the types of certain butterfly genera. The Ento-
mologist 66: 196-200.
. 1934a. Notes on nine genera of Holarctic butterflies. Stylops
3: 143-144.
. 1934b. New names for three genera of Rhopalocera. The
Entomologist 67: 77.
. 1935. Notes on seventeen genera of Rhopalocera. Stylops
4: 1-3.
. 1937. Changes in the genotypes of, or in the priority to be
accorded to, eleven genera of Lepidoptera, Rhopalocera, consequent
upon the determination of the dates of publication of the entomologi-
cal works of Jacob Hiibner. Stylops 6: 149-153.
. 1939. Notes on the generic nomenclature of the Lepidoptera
Rhopalocera I. Stylops 8: 133-138.
. 1943. Notes on the generic nomenclature of the Lepidoptera
Rhopalocera II. Stylops 12: 23-30.
. 1964a. Selection of type species for three nominal genera of
the family Satyridae. Annotationes Lep. (4): 120-121.
1964b. Introduction, as replacements for names invalid under
the Law of Homonymy, of names for three genera in the family Satyri-
dae. Annotationes Lep. (4): 137-138.
mem. amer. ent. soc, 24
158 THE SATYRIDAE
1964c. Fifteen nominal genera in the butterflies based on mis-
identified type-species, the names of which have been submitted to
the International Commission on Zoological Nomenclature for the des-
ignation under Article 70(a) of type-species in harmony with the in-
tention of the author of the generic name in question. Annotationes
Lep. (5): 157-171.
in press. Generic names of the butterflies. London, Trustees of
the British Mus.
Herrera, J. 1965. Etcheverrius y Palmaris nuevos generos de Satyridae
Andinos (Lepidoptera). Publ. Cent. Estud. Ent. (7): 57-73; ill.
. 1966. Quilaphoethosus, Chillanella y Haywardella nuevos
generos de Satyridae Andinos (Lepidoptera). Publ. Cent. Estud. Ent.
(8): 69-72; ill.
Herrera, J. and M. Etcheverry 1965. Stuardosatyrus nuevo genero de
Satyridae y revalidacion de la especie williamsianus Butler, 1868. Publ.
Cent. Estud. Ent. (7) : 74-77; ill.
Herrich-Schaffer, G. A. W. 1844 (1843-1856). Systematische Bearbei-
tung der Schmetterlinge von Europa, zeugleich als Text, Revision und
Supplement zu J. Hiibners Sammlung Europaischer Schmetterlinge. Re-
gensburg, Priv. publ.: 128 pis.
. 1864. Prodromus systematis Lepidopterorum. Corresp.-Blatt.
Zool.-Mineral. Verein. Regensburg 18: 1-52.
. 1865. Lepidopterorum index systematicus. Corresp.-Blatt
Zool.-Mineral. Verein. Regensburg 19: 63-76, 84-92, 100-108.
Hesse, R., W. C. Allee and K. P. Schmidt 1951. Ecological animal ge-
ography, 2nd ed. New York, John Wiley and Sons: xiii + 715 pp.; ill.
Hewitson, W. C. 1856-1876. Illustrations of new species of exotic butter-
flies. London, Van Voorst: 5 vols.
. 1865. A monograph of the genus Yphthima; with descriptions of
two new genera of diurnal Lepidoptera. Trans. Ent. Soc. London
(3)2: 281-293.
. 1877. Equatorial Lepidoptera. Part 5. London, Priv. publ.:
92 pp.
Holland, W. J. 1887. Notes upon a small collection of Rhopalocera made by
Rev. B. C. Henry in the island of Hainan, together with descriptions
of some apparently new species. Trans. American Ent. Soc. 14:
1 11-124.
. 1898. The butterfly book. New York, Doubleday, Page and
Co.: 382 pp.; ill.
Houlbert, C. 1922. Contribution a Fetude des Melanargiinae de Chine et
de Siberie. Etudes Lep. Comp. 19(2): 1-163; ill.
Hubner, J. 1806-1838. Sammlung Exotischer Schmetterlinge. Augsburg,
Priv. publ.: 4 vols.
. 1816-1826. Verzeichniss bekannter Schmettlinge. Augsburg,
Priv. publ.: 431 pp.
LEE D. MILLER 159
. 1818-1825. Zutrage zur Sammlung Exotischer Schmetterlinge,
I. Augsburg, Priv. publ.: 304 pp.
Huxley, J. (ed.), 1940. The new systematics. Oxford, The Clarendon
Press: 380 pp.
International Trust for Zoological Nomenclature 1961. International
Code of Zoological Nomenclature. London, Internatl. Trust Zool. No-
men.: xix + 176 pp.
Jordan, K. 1924. On Hypocysta and some allied genera of Satyrinae (Lep.
Rhop.) from New Guinea and the Solomon Islands. Novitates Zool.
31: 279-297; ill.
Just, T. 1948. Gymnosperms and the origin of the angiosperms. Bot. Gaz.
110(1): 91-103.
Karsch, F. 1893. Tagfalter-Lepidoptera Rhopalocera - von Adeli. Ber-
liner Ent. Zeits. 38: 167-266; ill.
. 1894. Eine neue ostafrikanische Lepidopteren-Gattung und -Art
aus der Familie der Satyriden. Ent. Nachr. 20: 190-192.
Kirby, W. 1837. Insects, in Richardson, J., Fauna Boreali-Americana: or
the zoology of the northern parts of British America. Part 4. London:
xxxix + 325 pp.; ill.
Kirby, W. F. 1871a. A synonymic catalogue of diurnal Lepidoptera. Lon-
don, Van Voorst: i-vii + 690 pp.; ill.
. 1871b. [Note]. Zool. Rec. [1871]: 363.
. 1877. A synonymic catalogue of diurnal Lepidoptera, Supple-
ment. London, Van Voorst: i-vii + pp. 691-883.
1894. A handbook to the order Lepidoptera. Part 1. But-
terflies, vol. 1. in Allen's Nat. Libr. Handbook. London: lxxiv +
261 pp.
. 1908 (1908-1912). Zutrage zur Sammlung exotischer Schmet-
terlinge. Neue deutsche facsimile ausgabe. Brussels, P. Wytsman:
i-iv + 1-101; ill.
Latreille, P. A. 1810. Considerations generales sur l'ordre natural des
animaux. . . crustaces. . . arachnides. . . insectes. . . Paris: 444 pp.
Leech, J. H. 1892. Butterflies of China. Japan and Corea, I. London, Priv.
publ.: 286 pp.
deLesse, H. 1951. Divisions generiques et subgeneriques des anciens genres
Satyrus et Eumenis (sensu lato). Rev. franc. Lep. 13: 39-42.
. 1952. Revision des anciens genres Pararge (S. L.) et Maniola
(=Epinephele (auct.)). Ann. Soc. Ent. France 121: 61-76.
von Linne, C. 1758. Systema naturae. . . regnum animale, ed. 10. Stock-
holm: 824 pp.
Mabille, P. 1880. [Note on Malgache butterflies]. Bull. Soc. Ent. France
(5)9: clxxiii-clxxiv.
. 1885-1887. Histoire naturelle des Lepidopteres. in Grandidier,
A., Hist. Phys. Nat. et Polit. Madagascar. Paris, L'Impremerie Na-
tionale: 364 pp.
MEM. AMER. ENT. SOC, 24
160 THE SATYRIDAE
Mathew, G. A. 1886. Descriptions of some new species of Rhopalocera
from the Solomon Islands. Proc. Zool. Soc. London, [1886]: 343-
350; ill.
Matthew, W. D. 1915. Climate and evolution. Ann. New York Acad.
Sci. 24: 171-318; ill.
Mayr, E. (ed.), 1952. The problem of land connections across the South
Atlantic, with special reference to the Mesozoic. Bull. American Mus.
Nat. Hist., 99(3): 85-258; ill.
. 1965. Numerical phenetics and taxonomic theory. Syst. Zool.
14(2): 73-97.
Meigen, J. W. 1829-1832. Systematische Beschreibung der Europaischen
Schmetterlinge. . . Aachen and Leipzig: 2 vols.
Moore, F. 1857. in Moore, F., and T. Horsefeld (1857-1859). A cata-
logue o; ths lepidopterous i .sects ii the museum o. thj Honourable
East India Company. London, E. India Co.: v + 440 + 6 pp.
. 1865. On the lepidopterous insects of Bengal. Proc. Zool. Soc.
London [1865]: 755-823; ill.
1878. A list of the lepidopterous insects collected by Mr.
Ossian Limborg in upper Tenasserim, with descriptions of new species.
Proc. Zool. Soc. London [1878]: 821-859; ill.
. 1880. On the Asiatic Lepidoptera referred to the genus Myca-
lesis; with descriptions of new genera and species. Trans. Ent. Soc.
London 28: 155-177.
. 1880-1881. Lepidoptera of Ceylon, 1. London, L. Reeve and
Co.: iii-xii + 190 pp.; ill.
. 1881. Descriptions of new Asiatic diurnal Lepidoptera. Trans.
Ent. Soc. London 29: 305-313.
. 1890-1893. Lepidoptera Indica. London, L. Reeve and Co.:
2 vols.: ill.
Muschamp, T. A. H. 1915a. The Ci-deviant genus Epinephele. Ent. Rec.
27: 152-156.
. 1915b. The genus Coenonympha. Mitt. Ent. Zurich 1: 20.
deNiceville, L. 1891. On new and little-known butterflies from the Indo-
Malayan region. Jour. Bombay Nat. Hist. Soc. 6: 341-398; ill.
. 1898. On new and little-known butterflies from the Indo-
Malayan, Austro-Malayan and Australian regions. Jour. Bombay Nat.
Hist. Soc. 12: 131-161; ill.
Novicky, S. 1923. Description d'un nouveau genre de Satyridae (Lepid.).
Bull. Sect. Ent. Mus. Nat. Prague 1: 60-61.
Oberthur, C. 1879. Catalogue raisonne des Papilionidae de la collection
de Ch. Oberthur a Rennes. Etudes. Ent. 4: 117 pp.
. 1916. Faune des Lepidopteres de Barbarie (partie ii). Etudes
Lep. Comp. 11: 179-428; ill.
LEE D. MILLER 161
Oberthur, C. and C. Houlbert, 1922. Quelques oues nouvelles sur la sys-
tematique des Melanargia (Lepidopteres: Satyridae). C. R. Acad.
Sci. Paris 174: 190-192.
Overlaet, F. G. 1955. Danaidae, Satyridae, Nymphalidae, Acraeidae.
Explor. Pare. nat. Upemba, Miss, de Witte (fasc. 27): 1-106: ill.
Prout, L. B. 1901. Staudinger and Rebel's Catalogue. Ent. Rec. 13: 230-
233.
Riley, N. D. 1922. The Rhopalocera of the Mt. Everest 1921 Expedition.
Trans. Ent. Soc. London 70: 461-483; ill.
Rober, J. 1892. See Schatz, E., and J. Rober, 1892.
Rothschild, W. and K. Jordan 1905. On some new Lepidoptera discovered
by A. S. Meek in British New Guinea. Novitates Zool. 12: 448-
478; ill.
Schatz, E. and J. Rober 1892. Die Familien und Gattungen der Tagf alter.
in Staudinger, O.. and E. Schatz, Exotische Schmetterlinge 2. Furth
(Bayern), G. Lowensohn: 284 pp.; ill.
Schrank, F. von P. von 1801 (1798-1803). Fauna Boica. Nurnberg: vol.
2 of 3 vols.
Schuchert, G. 1935. Historic geology of the Antillean-Caribbean region
or lands bordering the Gulf of Mexico and the Caribbean Sea. New
York, John Wiley and Sons: xxvi + 811 pp.: ill.
Schwanwitsch, B. N. 1924. On the ground-plan of wing-pattern in nympha-
lids and certain other families of the rhopalocerous Lepidoptera. Proc.
Zool. Soc. London [1924]: 509-528; ill.
. 1925. On a remarkable dislocation of the components of the
wing-pattern in the satyrid genus Pierella. The Entomologist 58: 266-
269: ill.
1931. Evolution of the wing-pattern in palearctic Satyridae.
II. Genus Melanargia. Zeits. Morphol. Oekol. Tiere 21: 316-408; ill.
Scopoli, G. A. 1777. Introductio ad historiam naturalem. . . Prague, Priv.
publ.: x + 506 pp.
Scudder, S. H. 1872. A systematic revision of some of the North American
butterflies; with brief notes on those known to occur in Essex Co.,
Mass. 4th Ann. Rept. Peabody Acad. Sci. [1871]: 24-83.
. 1875a. Historical sketch of the generic names proposed for
butterflies. Proc. American Acad. Sci. (2)10: 91-293.
. 1875b. Synonymic list of the butterflies of North America,
north of Mexico. Part 1, Nymphales. Bull. Buffalo Soc. Nat. Sci.
2: 233-269.
Sharpe, E. M. 1894. List of butterflies collected by Captain J. W. Pringle,
R. E., on the march from Teita to Uganda, in British East Africa. Proc.
Zool. Soc. London [1894]: 334-353; ill.
MEM. amer. ent. soc, 24
162 THE SATYRIDAE
Simpson, G. G. 1952. Probabilities of dispersal in geologic time, in Mayr,
E., 1952: 163-176.
. 1953. Evolution and geography. Eugene, Oregon (Condon
Lectures), Oregon State Syst. Higher Ed.: 1-64; ill.
. 1961a. Principles of animal taxonomy. New York, Columbia
Univ. Press: xii + 247 pp.; ill.
1961b. Historical zoogeography of Australian mammals.
Evol. 15(4): 431-446.
Sodoffsky, C. H. W. 1837. Etymologische Untersuchungen iiber die Gat-
tungsnamen der Schmetterlinge. Bull. Soc. Imp. Nat. Moscou 10(6):
76-91.
Sokal, R. R. and P. H. A. Sneath. 1963. Principles of numerical taxonomy.
San Francisco and London, W. H. Freeman and Co.: vii-xvi + 359
pp.; ill.
van Son, G. 1955. The butterflies of southern Africa. Part II. Nympha-
lidae: Danainae and Satyrinae. Pretoria, Transvaal Mus. Mem. (8):
v-ix +166 pp.; ill.
. 1958. A new African genus of the subfamily Satyrinae. Lep.
News 12: 6.
Staudinger, O. 1884-1888. in Staudinger, O., and E. Schatz, Exotische Tag-
falter. Exotische Schmetterlinge, 1. Furth (Bayern), G. Lowensohn:
iv + 334 pp.; ill.
. 1897. Neue sudamerikanische Tagfalter. Iris 10: 123-151.
Stichel, H. 1902. Aufteilung der Gattung Opsiphanes Westw., Beschreibung
neuer Brassoliden und synonymische Notizen. Berliner Ent. Zeits. 46:
487-524.
Strand, E. 1912. Zwei neue Satyriden von Roraima. Fauna Exot., Frank-
furt 2: 43-44, 47-48.
. 1942. Miscellanea nomenclatorica zoologica et palaeontologica.
X. Folia Zool. Hydrobiol. 11(2): 386-402.
Swainson, W. 1840. On the history and natural arrangement of insects.
London, Lardner's Cabinet Ecycl., [1-6]: 1-406.
Swinhoe, C. 1908. New eastern Lepidoptera. Ann. Mag. Nat. Hist. (8)1:
60-68.
Thieme, O. 1905. Monographic der Gattung Pedaliodes Butl. (Lepidoptera
Rhopalocera. Satyridae). Berliner Ent. Zeits. 50: 43-141.
. 1906. Monographische Bearbeitung der Gattungen Lasiophila
Felder, Daedalma Hew., Catargynnis Rober, Oxeoschistus Butl., Prono-
phila Westw., Corades Doubl. Hew. (Lepidoptera Rhopalocera. Saty-
ridae). Mit Begriindung neuer Gattungen und einer Anzahl Neubesch-
reibungen. Berliner Ent. Zeits. 51: 101-233; ill.
Tutt, I. W. 1896. British butterflies, being a popular handbook for young
students and collectors. London: 476 pp.; ill.
Verity, R. 1920. Contributo alle ricerche sulla variazione e la distribuzione
LEE D. MILLER 163
dei Lepidotteri in Italia: Zygaenides, Grypocera e Rhopalocera del Mas-
siccio delle Mainardi. Boll. Lab. Ziol. Portici 14: 33-61.
1953. Le farfalle diurne d'ltalia. Volume quinto. Divisione
Papilionoidea. Sezione Nymphalina: famiglia Satyridae. Florence,
Italy, Casa editrice Marzocco: xix + 354 pp.; ill.
Wallengren, H. D. J. 1853. Skandinaviens Dagfjarilar — Lepidoptera
Scandinaviae Rhopalocera, deposita et descripta. Malmo. Sweden: xx
+ 280 pp.
. 1857. Kaffernlandets Dag-Fjarilar, insamlade aren 1838-1845
af J. A. Wahlberg. K. Svenska Vet.-Akad. Handl. 2(4) : 1-55.
. 1858. Nya Fjaril-slagten. K. Svenska Vet.-Akad. Handl. 15:
75-84, 135-142.
Warren, B. C. S. 1936. Monograph of the genus Erebia. London, Trustees
of the British Mus. (Nat. Hist.): v-vii + 407 pp.: ill.
. 1947. Some principles of classification in Lepidoptera with spe-
cial reference to the butterflies. The Entomologist 80: 208-217. 235-
241.
Waterhouse, G. A. and G. Lyell. 1914. The butterflies of Australia. . .
Sydney, Angus and Robertson: iv-vi + 239 pp.; ill.
Watkins, H. T. G. 1925. New Callerebias (Lepidoptera. Satyridae). Ann.
Mag. Nat. Hist. (9)16: 233-237.
. 1928. New satyrid butterflies. Ann. Mag. Nat. Hist. (10)1:
615-618.
Westwood, J.O. 1841. //; Humphreys, H. N., and J. O. Westwood, British
butterflies and their transformations, ed. 1. London: xii + 138 pp.; ill.
Weymer, G. 1890. in Weymer, G., and P. Maassen, Lepidopteren gesam-
melt auf einer Reise durch Colombia, Ecuador, Peru, Brasilien, Argen-
tinien und Bolivien in den Jahren 1868-1877 von Alphons Stiibel. Ber-
lin, Priv. publ.: vi + 182 pp.
. 1909. Eine neue Gattung und zwei neue Arten aus der Familie
der Satyriden. Ent. Zeits. Stuttgart 23: 163-165.
1910-1912. Satyridae. in Seitz. A., Die Grossschmetterlinge
der Erde, vol. 5 (Die Amerikanische Tagf alter). Stuttgart, Alfred Ker-
nan Verlag: pp. 173-283; ill.
Wood-mason, J. 1880. Description of Parantirrhoea Marshallii, the type of
a new genus and species of rhopalocerous Lepidoptera from South India.
Jour. Asiatic Soc. Bengal 49(2): 248-250.
Zeller, P. C. 1850. Zwei neue Tagf alter. Stettiner Ent. Zeitg. 11: 308-313.
MEM. AMER. ent. soc, 24
164
THE SATYRIDAE
Index to Taxa
The numbers in bold face type indicate that the taxon is figured
on that page. Those names preceded by a $ mark are preoccupied
names.
abdelkader Pierret, 123
abeona Donovan, 81
abretia Capronnier, 92
achanta Donovan, 79, 81
acraeida Butler, 118
Acrophtalmia Felder & Felder, 67, 68, 69
" Acrophthalmia" Felder & Felder, 69
Acropolis Hemming, 69
actaea Linne, 121, 123
actoriaena Linne, 32-34
Ad mi ratio Hemming, 58, 62, 63
adolphei Guerin, 63
adoptiva Weymer, 31, 32
aega Fabricius, 132
aegeria Linne, 48, 50
aello Hiibner, 124
Aeropetes Billberg, 43, 44, 46. 47, 48, 52
Aeropetes-series, 40, 41, 42. 43. 47, 72,
137, 139, 151, 152
aethiops Esper, 105
Agapetes Billberg, 126
Agapetinae, 124
agondas Boisduval, 56, 57
Agrusia Moore, 56
albinotata Butler, 94
albonotata Godman, 117
albopunctata Weymer, 117
alini Bang-Haas, 103, 104
alope Fabricius, 101
A Itopedaliodes Forster, 1 1 4
Amathusia Fabricius, 132
Amathusiidae, 131
ambiorix Wallengren, 117
Amecera Butler, 50
Amechania Hewitson, 51, 52, 53
Amphidecta Butler, 110, 111, 113, 114
amphirhoe Hiibner, 27
Anadebis Butler, 50
anaxias Hewitson, 63
anaxioides Marshall & deNiceville, 63
Anchiphlebia Butler, 31
andromacha Hubner, 47
"andromeda Fabricius", 22
anthe Ochsenheimer, 123
antipodum Doubleday, 79, 80
Antirrhea Hubner, 9, 30, 31, 32
Antirrhini, 26, 28, 29-32, 34, 35, 36, 37,
133, 136, 138, 144, 145, 146
antonia Staudinger, 114
Antopedaliodes Forster, 114
Apaturina Herrich-Schaffer, 19
Aphantopus Wallengren, 95, 96, 97, 98
Aphysoneura Karsch, 43, 47, 48, 151
aramis Hewitson, 62
Aranda Fruhstorfer, 49
arcesilaus Cramer, 31
archaea Hubner, 31, 32
Archeuptychia Forster, 92
Archondesa Moore, 47
areolatus Smith & Abbot, 94
aretliusa Esper, 123
Arethusana deLesse, 123
XArge Hubner, 126
Arge Schrank, 126
arge Sulzer, 126
Argeformia Verity, 124, 126
argent ens Blanchard, 110, 112, 117
Argestina Riley, 87
Argus Bohadsch, 49
% Argus Scopoli, 49
Argynnina Butler, 75, 77, 80
Argyreuptychia Forster, 93
Argyronympha Mathew, 77, 80
Argyrophenga Doubleday, 74, 75, 78, 79,
80
Argyrophorus Blanchard, 109, 110, 112.
113, 117
armandi Oberthur, 123
armilla Butler, 94
Arpidea Duncan, 31
artemis Felder & Felder, 68, 69
ashna Hewitson, 94
asiatica Oberthur & Houlbert, 126
asochis Hewitson, 63
Atercoloratus Bang-Haas, 102, 103
atreus Kollar, 24
Auca Hayward, 110, 117
Aulocera Butler, 123
LEE D. MILLER
165
"aurora Felder", 22
automedon Cramer, 27
autonoe Cramer, 94
autonoe Esper, 123
buladeva Moore, 47
baldus Fabricius, 85
banghaasi Weymer, 94
bathseba Fabricius, 101
bega Westwood, 112, 117
Berberia deLesse, 123
bhadra Moore, 49
Bia Hiibner, 29, 30, 32-34, 136
Bicyclus Kirby, 58. 62, 63
Biina, 3, 4, 28, 32
Biinae, 3, 4, 8, 9, 11, 12, 13, 16, 17, 18,
19, 23, 28-36, 39, 72, 132, 133, 135,
136, 140, 144, 146, 147, 148
Biini. 10, 28, 29, 30, 32-34, 133, 144,
145, 146
Blanaida Kirby, 49
blanda Moschler, 93
Blelogona Felder & Felder, 40, 57, 58,
59, 60, 61
Boeberia Prout, 86, 87
bolanica Marshall, 85
Bolboneura Godman & Salvin, 16
brahminus Blanchard, 123
Brassolidae, 4, 23
Brassolides, 2, 23
Brassolina, 3, 4
Brassolinae, 2, 3, 4, 8, 9, 11, 13, 16, 17,
18, 19, 23-28, 29, 72, 133, 136, 140,
144, 145, 146
Brassolini, 23-28, 133, 144, 145
Brassolis Fabricius, 24, 25, 26, 27
Brassolis-series, 25, 26, 27
Brintesia Fruhstorfer, 121, 123
briseis Linne, 123
Bruasa Moore, 54, 56
butleri Fereday, 79, 80
byses Godart, 93
Caerois Hiibner, 8. 30, 31, 32
caerulea Butler, 92
Caeruleuptychia Forster, 92
Caliginae, 3, 4, 23
Caligo Hiibner, 23, 24, 25, 26, 27
Caligo-series, 25, 27
Calislo Hiibner, 113. 114
Callarge Leech, 51, 52, 53
Callerebia Butler, 82, 86, 87
Callerebia-series, 83, 87, 102, 141. 147,
148
Callicore Hiibner, 16
Callinaga Moore, 19
callipteris Butler, 47
Callitaera Butler, 22, 134
Callyphthima Butler, 86
Calysisme Moore, 61
Capronnieria Forster, 92
cassiae Linne, 24
cassina Butler, 108
Cassionympha vanSon, 86
cassiope Cramer, 27
cassiope Fabricius, 105
cassius Godart, 86
Cassus vanSon, 106, 107, 108
cassus Linne, 107, 108
Catargynnis Rober, 112, 114
Catoblepia Stichel, 25, 27
cecilia Vallantin, 101
Celebina Fruhstorfer, 61
Cepheuptychia Forster, 92
cephus Butler, 92
Cercyonis Scudder, 99, 100. 101, 120,
145
ceto Hiibner, 104
Charma Doherty, 47
Chazara Moore, 123
Cheimas Thieme. 112, 114
clwiiui Guerin, 85
"chiliensis C. et R. Felder". 118
chiliensis Guerin, 118
Chillanella Herrera, 118
Chionobas Boisduval, 120, 124
Chloreuptychia Forster, 92
clitoris Cramer, 92
Chonala Moore, 49
Choranesa Moore, 47
chorinaeus Fabricius, 31, 32
Chortobius [Dunning & Pickard], 96, 97,
98
christophi Leech, 49
circe Fabricius, 121, 123
cirta Felder & Felder, 1 1 6
Cissia Doubleday, 91, 92
MEM. AMER. ENT. SOC, 24
166
THE SATYRIDAE
Cithaerias Hiibner, 20, 21, 22
clarissa Cramer, 92
clio Weymer, 94
Clothilda Blanchard, 133
cluena Drury, 92
clyte Hiibner, 108
clytus Linne, 107, 108
Coelites Westwood & Hewitson, 64, 65,
66, 67, 72
Coenonympha Hiibner, 96, 97, 98, 142,
145
Coenonymphini, 9, 10, 11, 70, 73, 74, 75,
81, 89, 95-98, 99, 142, 144, 146, 147
Coenyropsis vanSon, 86
Coeruleotaygetis Forster, 93
Colaenis Hiibner, 19
constantia Cramer, 35, 36
Corades Doubleday, 112, 114
corderoi Dognin, 114
Corderopedaliodes Forster, 114
Cosmosatyrus Felder & Felder, 118, 127
Crebeta Moore, 44, 48, 150
creola Skinner, 145
crisia Geyer, 68, 69
Culapa Moore, 59, 60, 61
cyclopina Staudinger, 115, 118
cyllastros Westwood & Hewitson, 24, 28
Cyllo Boisduval, 36
Cyllogenes Butler, 30, 35, 36
Cyllopsis R. Felder, 90, 91, 93
cymela Cramer, 87, 90, 91, 94, 147
Daedalma Hewitson, 109, 112, 116
Dalapa Moore, 58, 59, 61
Dallacha Moore, 82, 83, 85
damaris Doubleday, 50
Danaidae, 15, 19, 55, 130, 134
Danaus Latreille, 130
% Dasyomma Felder & Felder, 61
Dasyomma Macquart, 61
Dasyophthalma Westwood, 24, 25, 26, 27
Dasyophthalma-series, 25, 27
Davidina Oberthur, 119, 120, 121, 122.
123
Debis Doubleday & Hewitson, 47
decorata Butler, 117
decorata Felder & Felder, 116
deidamia Eversmann, 48, 50
dendrophilus Trimen, 47, 48
dexamenus Hewitson, 61
diademoides Moore, 50
Dichothyris Karsch, 58, 62, 63
Didonis Hiibner, 56
digna Marshall, 123
Dingana vanSon, 106, 108
dingana Trimen, 108
dinias Hewitson, 112, 116
Dionana Moore, 47
Dioriste Thieme, 111, 116
Dira Hiibner, 106, 107, 108
Dirini, 9, 10, 12, 41, 71, 72, 73, 102,
105-108, 119, 124, 139, 143, 144, 146.
150-151, 152
Dodonidia Butler, 74, 75, 78, 79, 80
doraete Hewitson, 116
dor us Esper, 98
drepana Westwood, 62, 63
Drucina Butler, 110, 111, 116
dntsia Cramer, 61
drusillodes Oberthur, 61
Drusillopsis Fruhstorfer, 61
dryas Scopoli, 123
Dubierebia Muschamp, 98
Dulcedo d'Almeida, 20, 22
dumetorum Oberthur, 48, 50
duplex Butler, 87
duponcheli Guerin, 63
Dyctis Boisduval, 55, 56, 57
Dynastor Westwood & Hewitson, 25, 26,
27
dynsate Hewitson, 47
Elina Blanchard, 109, 115, 117
Elina-series, 108, 110, 117
Elymnias Hiibner, 54, 56, 57, 132, 150
Elymniina, 3, 4, 38, 54
Elymniinae, 3, 4, 8, 9, 12, 13, 16, 17,
18, 19, 23, 26, 29, 38-63, 64, 67, 72,
135, 138, 139, 140, 144, 146, 147, 149.
150
Elymniini, 10, 11, 12, 13, 17, 18, 23, 26,
39, 40, 41, 42, 52, 54-57, 135, 137,
139, 144, 146, 150. 152
Elymniopsis Fruhstorfer, 54, 55, 56, 57
emma Staudinger, 116
Enodia Hiibner. 40, 41, 43, 46, 47
Enodiinae, 3, 4, 38, 40, 41
Enodiini, 41
t Enope Moore, 49
Enope Walker, 49
LEE D. MILLER
167
enyo Hewitson, 112, 114
Epigea Hiibner, 104
Epimede Houlbert, 126
epimenides Menetries, 50
Epinephele Hiibner, 101. 109
"Epinephila" auctt., 101
"Epinephile" auctt., 101
epiphron Knoch, 105
epistygne Hiibner, 103, 105
Erebia Dalman, 74, 78, 80, 102, 103,
104, 105, 108, 114, 142, 145
Erebiinae, 101
Erebiini, 9, 71, 73, 81, 95, 98, 101-105,
108, 119, 142, 144, 146, 147
erebioides Felder & Felder, 115, 118
Erebiola Fereday, 74, 78, 79, 80
t Erebonwrpha Elwes, 87
Erebomorpha Walker, 87
Eretris Thieme, 116
ericluho Butler, 93
Ericluhodes Forster, 93
Elites Westwood, 64, 65, 66
Eritinae, 4, 8, 9, 12, 16, 17, 19, 39, 64-
66, 67, 72, 135, 139, 140, 144. 146,
148, 149
Eritini, 64-66, 144
Erycinidia Rothschild & Jordan, 75. 76,
80
Eryphanis Boisduval, 25, 26, 27
esaca Westwood & Hewitson, 56
Etcheverrius Herrera, 118
Eteona Westwood, 109, 111, 116
Ethope Moore, 44, 50
Eumenis Hiibner, 123
euphemia Westwood & Hewitson, 76, 80
Euploea Fabricius, 55
Euploeamima Holland, 50
Euptychia Hiibner, 90, 91, 93
Euptychiini, 9, 70, 73, 74, 81, 87. 88,
89-95, 98, 99, 140, 141, 142, 144, 145,
146, 147, 148, 149
Euptychoides Forster, 93
euripides Weymer, 117
europa Fabricius, 45, 48
eurydice Linne, 46, 49, 145
eurylochus Hiibner, 24, 27
Eurytelidae, 38, 54
eurytus Fabricius, 94
evadne Cramer, 61
fagi Scopoli, 123
Faunis Hiibner, 132
Faimula Felder & Felder, 118
fergana Oberthiir & Houlbert, 126
francisca Stoll, 60, 61
fumata Butler, 95
fuscum Felder & Felder, 60, 61
galathea Linne. 125, 126
t Gallienia Oberthiir, 63
G are ris Moore, 61
"Geirocheilus" Holland, 116
Geitoneura Butler, 78, 79, 80
germainii Felder & Felder, 118
geticus Esper, 98
glacial is Moll, 124
Gnophodes Westwood, 36
Godartiana Forster, 93
Gorgo Hiibner, 102, 104
gotama Moore, 63
gracilis Rothschild & Jordan, 76, 80
Gyrocheilus Butler, 109, 110, 111. 112.
113, 116
gyrtone Berg, 1 17
Haetera Fabricius, 20, 21, 22, 134
Haeterinae, 3, 4, 7, 8, 9, 10, 11, 12. 13.
15, 16, 17, 18, 19-22, 23, 29, 72, 132.
133, 134, 136, 140, 144, 145, 146
Haeterini, 19, 20-22, 133, 144, 145
Halimede Oberthiir & Houlbert, 126
haiimede Menetries, 126
Hallelesis Condamin, 63
Hamadryopsis Oberthiir, 61
"Hanifa" auctt., 47
hanifa Nordmann, 123
Hanipha Moore. 47
Hariina Moore. 47
Harjesia Forster, 93
Harsiesis Fruhstorfer, 80
haydenii Edwards, 145
Haywardella Herrera, 116
Haywardina Aczel, 93
t Haywardina Forster, 93
hedemanni R. Felder, 91, 93
helmsii Butler, 79, 80
Hemadera Moore, 85
Henotesia Butler, 63
hercyna Hiibner, 36, 37
MEM. AMER. ENT. SOC, 24
168
THE SATYRIDAE
hermes Fabricius, 93
Hermeuptychia Forster, 93
Hermianax Fruhstorfer. 50
Hermias Fruhstorfer, 42, 44, 47
hermione Linne, 123
herse Cramer, 93
hesione Cramer, 62
hesione Sulzer, 94
Hetaerina, 3, 4, 19, 20
Heteronympha Wallengren, 75, 77, 81
Heteropsis Westwood, 58, 62, 63
hewitsonii Doumet, 62, 63
hilda Westwood, 50, 51
himachala Moore, 50
Hipio Hiibner, ,30, 35, 36
Hipparchia Fabricius, 119, 123
Hipparchiadae, 15, 69, 119
Hipparchioid.es Butler, 81
hippia Cramer, 87
hobartia Westwood & Hewitson, 80
Homoeonympha Felder & Felder, 109,
118
Houlbertia Oberthiir, 63
huebneri Felder, 123
huebneri Kirby, 85
humilis Felder & Felder, 118
hyagriva Moore, 85
Hyalodia Jordan, 75, 80
hygeia Hewitson, 80
hypaesia Hewitson, 22
"hyperanthus" auctt., 98
hyperantus Linne, 97, 98
hyperbius Linne, 86, 87
hypermnestra Linne, 56, 57
Hypocysta Westwood & Hewitson, 75,
76, 80
Hypocysta-series, 12, 74, 76-77, 78, 80,
81, 149, 150
Hypocystini, 7, 9, 11, 12, 57, 70. 72, 73.
74-81, 102, 105, 108, 110, 119, 124,
139, 140. 141, 144, 146, 149, 150,
151
Hyponephele Muschamp, 101
I data deLesse, 101
Idiomorphus Chaudoir, 63
i Idiomorphus Doumet, 63
t Idioneura Felder & Felder, 118
Idioneura Selys, 1 1 8
Idioneurula Strand, 110, 113, 115, 118
incerta Hewitson, 51, 53
Indalasa Moore. 61
inga Fruhstorfer, 61
inica Hewitson, 85
Ithomiidae, 9, 12, 14-15, 16, 19, 130,
131-132, 134
itonia Hewitson, 85
jalaurida deNiceville, 49
janardana Moore, 61
janira Linne, 101
J at ana Moore, 61
jurtina Linne, 100, 101
jynx Hiibner, 57
Kabanda Moore, 61
kalinda Moore, 87
Kallima Doubleday, 19
Kanetisa Moore, 123
Karanasa Moore. 119, 123, 148
Kerrata Moore, 47
Kirinia Moore, 4.3, 50
Kirrodesa Moore, 47
klugii Guerin, 79, 80
Kolasa Moore, 82, 85
Lachesis Oberthiir & Houlbert, 126
lachesis Hiibner, 126
lais Cramer, 57
lais Fabricius, 57
Lampides Hiibner, 130
Lamprolenis Godman & Salvin, 72, 74,
75, 76, 77, 80, 139
lanaris Butler, 47
languida Butler, 94
Lasiommata Westwood, 50
Lasiophila Felder & Felder, 110, 116
lathionella Westwood, 81
latipicta Fruhstorfer, 50
leaena Hewitson, 118
leda Gerstacker, 87
leda Leech, 126
leda Linne, .35. 36, 136, 150, 152
Ledargia Houlbert, 124, 126
lemur Schrank, 101
leonata Butler, 111, 116
leprea Hewitson, 81
Leptoneura Wallengren, 108
LEE D. MILLER
169
leptoneuroides Felder & Felder, 118
Lepiotes Scudder, 130
Lethe Hubner, 16, 41, 43, 44. 45, 47, 48
Le//je-series, 42, 43, 47-49, 137
Lethinae, 3, 4, 38, 40, 41
Lethini, 28, 34, 37, 38, 39, 40-51, 52. 54.
57, 64, 72, 74, 105, 135, 137, 138-
140, 144, 146. 147, 149, 151, 152
leucoglene Felder & Felder, 1 1 8
libye Linne, 94
Libythea Fabricius, 130
Libytheidae, 130
ligea Linne, 103. 104
Limenitinae, 52
Use Hemming. 56. 57
Loesa Moore, 61
Lohana Moore, 85
Lohora Moore, 61
Lopinga Moore, 44, 48, 50
lorquinii Felder & Felder, 46, 49
lowi Doubleday & Hewitson, 50
Loxerebia Watkins, 82, 87
Lycaena Fabricius, 74
Lycaenidae, 130
lycaon Rottemburg, 101
Lycoreinae, 133
Lyela Swinhoe. 95, 96, 98
Lymanopoda Westwood, 109, 118
Lymanopoda-scnes, 108, 110, 113, 114,
118, 127
lymessa Hewitson, 81
macmahoni Swinhoe, 98
madura Horsefield, 66
Magneuptychia Forster, 94
t Magula Fruhstorfer, 44, 49
Magula Scudder, 49
maianeas Hewitson, 63
malsara Moore, 49
mahahda Butler. 61
Manataria Kirby, 10, 18. 34. 36. 37, 38.
41, 146
Mandarinia Leech, 41, 43, 46, 50, 51,
72, 74, 139. 149
Mandarinia-stries. 42, 43, 50
Manerebia Staudinger, 110, 111, 115.
118
Maniola Schrank 2, 99. 100. 101, 109
Maniolidi, 98
Maniolinae, 69, 98
Maniolini. 9, 71, 73, 74, 81, 89, 98-
101, 109, 120, 142, 144, 146, 147
margaretae Elwes, 47
Market Hubner, 104
marshallii Wood-Mason, 36
Martanda Moore, 58, 59, 61
Mashuna vanSon, 83, 84, 85
mashuna Trimen, 84, 85
masoni Elwes, 49
Masoura Hemming, 58, 63
masoura Hewitson, 59, 63
inedus Fabricius, 60, 62
medus Schiffermiller, 104
Medusia Verity, 102, 104
megera Linne, 50
Megeuptychia Forster, 94
Megisto Hubner, 91, 94
Melampias Hubner, 82, 86, 87
Melampias-series, 83, 84, 86-87, 141.
151, 152
Melanargia Meigen, 4, 11, 16, 124, 125,
126
Melanargiinae, 124
Melanargiini, 5, 9, 10, 71, 73, 105. 119.
124-126, 143, 144, 146, 147
Melania Lamarck, 123
t Melania Sodoffsky, 123
Melanitini, 10, 18, 28, 29, 30, 34-36, 37,
39, 41, 135, 136, 138, 144, 146, 149,
152
Melanitis Fabricius, 16, 30, 35, 36, 150
Melynias Moore, 54, 55, 57
menander Drury, 22
Meneris Westwood, 47
mermeria Cramer, 91, 95
merope Fabricius, 81
Mestra Hubner, 11, 16
mestra Hewitson, 62
Mimadelias Moore, 54, 57
minerva Fabricius, 49
Minois Hubner, 99, 119, 120, 123
Mint ha vanSon, 108
mintha Geyer, 107, 108
mirabilis Staudinger, 118
mixturata Alpheraky, 95, 146
mnasicles Hewitson, 60, 61
mollina Hubner, 91, 92, 93
monachus Blanchard, 117
MEM. AMER. ENT. SOC, 24
70
THE SATYRIDAE
Moneuptychia Forster, 94
Monotrichtus Hampson, 61
monticolens Butler, 118
moorei Felder, 61
Morphidae, 2, 3, 14-15, 16, 19, 23, 131,
132, 134
Morpho Fabricius, 15, 132
Muscopedaliodes Forster, 1 1 6
muscosa Thieme, 116
Mycalesini, 11, 12, 13, 33, 39, 40, 52,
54, 57-63, 72, 74, 75, 128, 135, 138,
139, 144, 146, 147, 149, 150, 151, 152-
153
Mycalesis Hubner. 42, 46, 58, 60, 61,
63, 139
Mycalesis group, 67
mycalesis Felder & Felder, 61
mycalesoides Felder, 94
Mydosama Moore, 60, 61
Mygona Westwood, 116
mynois Hewitson, 61
myops Staudinger, 98
Myrtilus deNiceville, 61
mystes deNiceville, 61
Nadiria Moore, 85
napoleon Westwood & Hewitson, 27
narasingha Moore, 85
nareda Kollar, 85
Narope Westwood & Hewitson, 23, 24,
25, 26, 28
N a rope-scries, 25, 28
Nasapa Moore, 62
natalii Boisduval, 86
Nebdara Moore, 62
necys Godart, 93
Neita vanSon, 87
neita Wallengren, 87
Nelia Hayward, 109, 117
Nemetis Moore, 49
nemyroides Blanchard, 117
Neocoenyra Butler, 87
Neohipparchia deLesse, 123
Neomaenas Wallengren, 109, 117
Neomaniola Hayward, 117
Neominois Scudder, 119, 120, 121, 123
Neonympha Hubner, 69, 87, 91, 94
Neope Butler, 49
Neorina Westwood, 43, 50, 51
Neorina-series, 37, 39, 40, 42, 43, 50, 52,
138
Neosatyrus Wallengren, 117
nereis Drury, 21, 22
nesaea Linne, 57
Nesoxenica Waterhouse & Lyell, 77, 81
Ninguta Moore, 49
Nissanga Moore, 62
nitida Godman & Salvin, 76, 80
noma Thunberg, 122, 124
nothis Westwood & Hewitson, 66
Nymphalidae, 52, 130
Nymphaloidea, 7, 8, 19
Nytha Billberg, 123
obsoleta Westwood, 118
Ocaiis Westwood, 94
occitanica Esper, 126
Odonata, 131
oedipe Hubner, 98
oedippus Fabricius, 97, 98
Oeneis Hubner, 16, 119, 120, 122, 124,
142, 145
Oeneis-series, 73, 120, 121, 122, 124,
143
opalina Butler, 87, 88
opalinus Staudinger, 114
Opoptera Aurivillius, 25, 27
Opsiphanes Westwood & Hewitson, 24,
25, 26, 27
Oreas Hubner, 22
Oreina Westwood, 105
Oreixenica Waterhouse & Lyell, 81
oreseis Hewitson, 63
Oressinoma Westwood, 90, 92, 93, 94
Orinoma Gray, 43, 50
ornata Rothschild & Jordan, 80
oroatis Hewitson, 61
Orsotriaena Wallengren, 40, 57, 58, 59,
60, 62, 139
Orthoptera, 131
Oxeoschistus Butler, 116
Pachama Moore, 62
Palaeonympha Butler, 74, 87-89, 146
pales Philippi, 117
Pal maris Herrera, 118
Pampasatyrus Hayward, 117
Pamperis Heimlich, 127-128
pamphanis Westwood & Hewitson, 27
pamphilus Linne, 97, 98
LEE D. MILLER
171
Panarche Thieme, 116
panda Boisduval, 84, 87
Pandima Moore, 83, 85
pandoea Hopffer, 63
Panyapedaliodes Forster, 1 1 6
panyasis Hewitson, 1 1 6
Papilio Linne, 2
Papilionoidea, 3
parado.xa Mabille, 62, 63
"Parage" auclt., 50
Paralasa Moore, 82, 83, 87
Paralethe vanSon, 9, 43, 44, 47, 48
Paramecera Butler, 90, 93, 94
Parantirrhoea Wood-Mason, 30, 34, 35.
36. 138
Parapedaliodes Forster, 116
"Pararga" auctt., 50
Pararge Hubner, 42, 43, 45, 48, 50
Para/ge-series, 39, 42, 43, 44, 46, 49-50,
137, 139, 147
Parataygetis Forster, 94
Paratisiphone Watkins, 81
Parce Oberthiir & Houlbert. 126
parce Staudinger, 126
pare pa Hewitson, 116
Pareuptychia Forster, 94
parmenio Boeber, 86, 87
parmeno Westwood, 36
Paroeneis Moore, 123
Paryphthimoides Forster, 94
Pasiphana deLesse, 101
passandava Ward, 63
Patala Moore, 49
patnia Moore, 62
patrobas Hewitson, 113, 116, 147
Pedal iodes Butler, 111, 113, 116
pegala Fabricius, 100, 101
pelopea Klug, 123
penanga Westwood & Hewitson, 56
penelea Cramer, 94
penelope Fabricius, 91, 93
Penetes Westwood & Hewitson, 25, 27
Penrosada Brown, 113, 118
Percnodaimon Butler, 74. 78, 79, 80, 81
periboea Fabricius, 128
periboea Godman & Salvin, 93
Periplysia Gerstacker, 83, 87
phaedra Linne, 123
phanias Hewitson, 116
phantoma Fassl. 22
phares Godart, 94
X Pharia Fruhstorfer, 69
Pharia Gray, 69
Pharneuptychia Forster, 94
t phegea Fabricius, 57
Pherepedaliodes Forster, 116
pheretiades Smith & Kirby, 116
phidia Linne, 123
Philareta Moore, 123
philomela Linne, 84, 85
pholoe Staudinger, 112, 114
Phorcis Hubner. 102, 103, 104, 105
phoronea Doubleday, 117
X Phryne Herrich-Schaffer, 98
Phryne Meigen, 98
phryne Pallas, 97, 98
Physcaeneura Wallengren, 83, 84, 87
physcoa Hewitson, 116
Physcon deNiceville, 63
Physcopedaliodes Forster, 116
piera Linne, 21, 22
Pierelta Herrich-Schaffer. 4, 20, 21. 22.
134
Pierellinae, 3, 4, 19, 20
Pieridopsis Rothschild & Jordan, 75, 76,
78, 80
pigmentaria Karsch, 47, 48
pignerator Butler, 111, 114
pimplea Erichson, 51, 52, 54
Pindis R. Felder, 90, 94
pireta Cramer, 21, 22
Placilla Moore, 49
Platypthima Rothschild & Jordan, 75,
77, 80
pluto Fereday, 79, 81
poaoeneis Heimlich. 127
poesia Hewitson, 111, 116
polita Hewitson, 22
poltys Prittwitz, 94
Polymastus Thieme, 116
polyxo Godman & Salvin, 117
portlandia Fabricius, 46, 47, 145
Posteuptychia Forster, 94
Posttaygetis Forster, 94
Praefaunula Forster, 94
Praepedaliodes Forster, 116
Praepronophila Forster, 116
pratorum Oberthiir, 87
MEM. AMER. ENT. SOC, 24
172
THE SATYRIDAE
Precis Hiibner, 130
pringlei Sharpe, 47
Proboscis Thieme, 113, 116
pronoe Esper, 105
Pronophila Doubleday & Hewitson, 109,
111, 112, 116
Pronophila-seiies, 10, 110, 114-117
pronophila Felder & Felder, 116-117
Pronophilinae, 3. 4, 108
Pronophilini, 7. 9, 10, 71, 73, 102, 105,
108-118, 119, 127, 140. 143, 144, 145,
146, 147, 148
propylea Hewitson, 113, 116
Pseudeuptychia Forster, 94
Pseudochazara deLesse, 123
Pseudodebis Forster, 94
Pseudohaetera Brown, 20, 22
% Pseudomaniola Rober, 114
Pseudomaniola Weymer, 114, 117
Pseudonympha Wallengren, 82, 87
Pseiidosteroma Weymer, 116-117
Pseudotergumia Agenjo, 123
Psyche Hiibner, 126
Pry chandra Felder & Felder, 34, 39, 41,
43, 44, 45, 46, 49, 138
puerta Hewitson, 116
pulchra Mathew, 80
pumilus Felder, 123
Punapedaliodes Forster, 117
pusilla Felder & Felder, 118
Putlia Moore, 47
Pyronia Hiibner, 100, 101
Quilaphoetlwsiis Herrera, 117
Ragadia Westwood, 9, 67, 68, 69
Ragadiina, 3, 4, 67
Ragadiinae, 3, 4, 8, 9, 12, 13, 15, 16, 17,
18, 39, 67-69, 72, 139, 141, 144, 146,
147, 148, 149
Ragadiini, 67-69, 144
Rangbia Moore, 45, 49
Rareuptychia Forster, 94
regalis Leech, 50, 51, 57, 147
Rhaphicera Butler, 45, 46, 50
Rliaphiceropsis Sharpe, 47
ridingsii Edwards, 121, 123, 145, 148
ruscinonensis Oberthiir & Houlbert, 126
rusina Godart, 24, 27
Sabatoga Staudinger, 1 1 8
Sadarga Moore, 63
safitza Hewitson, 61
sagitta Leech, 51, 53
saitis Hewitson, 77, 80
sallei Westwood & Hewitson, 27
Samanta Moore, 10, 18, 34, 39, 41, 49,
138
sambulos Hewitson, 62, 63
samio Doubleday & Hewitson, 47
samius Westwood, 110, 115, 118
Samundra Moore, 63
sanatana Moore, 61
Sarromia Westwood, 109, 118
Satoa Moore, 58, 63
satricus Westwood & Hewitson, 45, 50
saturnus Butler, 93
Satyrides, 2, 6, 15, 69, 119
Satyrina, 3, 4
satyrina Bates, 94
satyrina Butler, 45, 49
Satyrinae, 3, 4, 6, 7, 8, 9, 10, 12, 13,
16, 17, 18, 39, 41, 42, 57, 64, 67, 69-
126, 136, 139, 140-142, 144, 145, 146,
147, 148, 149, 150, 151, 152
Satyrini, 6. 7, 9, 10, 71, 73, 99, 105, 106,
108, 110, 119-124. 142, 143, 144, 145,
146, 147, 148, 149
Satyrini-section, 12, 73. 74, 75, 101, 105,
108, 110, 140, 141, 143, 145, 150
Satyrodes Scudder, 43, 44, 46, 49
Satyrotaygetis Forster, 94
Satyrus Latreille. 2, 119, 121, 123
Sarynw-series, 10, 120, 122, 123, 143,
148
scanda Kollar, 86, 87
scanda Moore, 45, 49
schrenkii Menetries, 49
Selenophanes Staudinger. 25. 27
servilaea Wallengren, 117
sesara Hewitson, 84, 85
Setodocis Billberg, 128
Sevanda Moore, 63
Sicca Verity, 98
sicelis Hewitson, 47
sidonis Hewitson, 49
sihala Moore, 47
similis Butler, 95
Sim plica Verity, 105
Sinarista Weymer. 29, 30, 31. 32
LEE D. MILLER
173
Sinchula Moore, 49
% Smithia Mabille, 63
Smithia Milne-Edwards & Haime, 63
sophorae Linne, 24. 26. 27
soter Butler, 94
Spinantenna Hayward, 110, 114. 115, 117
Splendeuptychia Forster, 94
squamistriga R. Felder. 94
statilinus Hufnagel, 123
stelligera Butler. 1 1 8
Steremnia Thieme, 1 17
Steroma Westwood, 109, 112, 113, 114,
117
Steromapedaliodes Forster, 117
Stibonwrpha Butler, 117
Strabena Mabille, 82, 83. 87
Stuardosatyrus Herrera & Etcheverry,
117
Stygionympha vanSon, 87
stygne Ochsenheimer. 104
Stygnolepis Strand, 118
t Stygnus Felder & Felder, 1 1 8
Stygnus Perty, 118
sudra Felder, 61
sura Westwood, 49
suradeva Moore, 36
Suralaya Moore, 63
syme Hiibner, 27
Syngea Hiibner, 105
Taenaris Hiibner, 19, 132
tagala Felder, 62
tamatavae Boisduval, 87
Tanaoptera Billberg, 48
Tansima Moore, 42, 45. 49
Tarsocera Butler, 106, 108
Tatinga Moore, 50
tauropolis Westwood. 1 1 6
Taygetina Forster, 94
Taygelis Hiibner. 90, 91. 92. 95
Telinga Moore, 63
Tellervini, 132
tena Hewitson, 1 14
tenuisquamosa Joicey & Talbot. 80
Tetraphlebia Felder & Felder, 118
thalia Leech, 69
t thelebe Doubleday, 116
thelebe Doubleday & Hewitson, 111, 116
Theope Doubleday & Hewitson, 50
X Theope Moore, 50
thibetanus Oberthiir, 50
Thiemeia Weymer, 1 1 7
thione Berg, 1 1 7
Thympia Moore. 82, 85
tircis Stoll, 98
Tisiphone Hiibner, 81
tisiphone Boisduval, 116
tithone Hiibner, 101
tithonus Linne, 100. 101
Tithoreini, 132
tomasia Butler, 31
Torynesis Butler, 10, 106, 107. 108
Triariia Verity, 105
triarius dePrunner, 105
tricordata Hewitson, 116
trimacula Leech, 47
Triphysa Zeller, 95, 96, 97, 98
tristigmata Elwes, 47
tristis Guerin, 115. 117
Triteleuta Strand. 30, 3 1
Trophonina Rober, 118
Truucaefalcia Verity, 105
tulbaghia Linne. 47, 48
typhla Westwood, 92, 93, 94
valentina Cramer, 94
Vanessa Fabricius, 130
vanessoides Blanchard, 115. 117
Vareuplychia Forster, 95
vasudeva Moore, 57
verma Kollar, 47
vigilans Trimen, 87
Virapa Moore, 63
virgilia Cramer, 95
virgo Rothschild & Jordan, 76, 80
viridicans Weymer, 95
vitellia Cramer, 56
waltoni Elwes. 87
wardii Butler, Callyphthima, 86
wardii Butler, Henotesia, 63
weidemeyeri Edwards, 52
Weymerana Forster. 95
williamsianus Butler, 117
Xenica Westwood, 75, 78, 79, 81
Xenica-scries, 74. 76, 77, 78. 80-81, 108,
150
MEM. AMER. ENT. SOC, 24
174
THE SATYRIDAE
Xeniconympha Novicky, 81
xicaque, Reakirt, 93, 94
Xois Hewitson, 82, 84, 85, 150
yama Moore, 49
"Yphthima" auctt., 85
yphthima Felder, 95
Yphthimoides Forster, 95
Ypthima Hiibner, 82, 84. 85
Ypthima group, 67
Ypthimaseties, 82, 83, 84, 141
Ypthimini, 9, 10, 70, 73, 74, 75, 81-87,
88, 89, 95, 99, 101, 102, 141, 144, 145,
146, 147, 149, 150, 151, 152
Ypthimini-section, 12, 73, 74, 75, 81, 90,
95, 99, 102, 140, 141, 145
Ypthimomorpha vanSon, 85
Ypthimorpha Overlaet, 85
Zabimia Hewitson, 118
zangris Hiibner, 114
Zethera C. Felder, 51, 52, 53, 54
Zetherini, 10, 39, 40, 41, 52-54, 135, 139,
144, 146, 147
zigomala Hewitson, 118
Zipaetis Hewitson, 75, 77, 78, 80
Zischkaia Forster, 95
Zophoessa Westwood, 43, 49
Mi
An ii
spective subje
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Keys for the
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r than thu
of the R (Colcoptera).
kiel Rivnay. 1929. $2
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